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A classification of the Lepidoptera 
based on characters of the pupa 



Edna Mosher 



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Bulletin of the Illinois State Laboratory of Natural History. 
Vol. XII, Art. II. 



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A CLASSIFICATION OF THE LEPIDOPTERA 
BASED ON CHARACTERS OF THE PUPA 



BY 



EDNA MOSHER 

B. S. A., Cornell Univerfity, 1908 
M. S., University oi Ulinoif, 1913 



. THESIS 

Submitted in Partial Fulfilment oi the Requirements for tke 

Degree oi 

DOCTOR OF PHILOSOPHY 
IN ENTOMOLOGY 

IN 

THE GRADUATE SCHOOL 

OF THE 

UNIVERSITY OF ILLINOIS 
I9I5 



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CONTENTS 



PAGE 

Introduction 17 

Changes preceding pupation 18 

External morphology 22 

Classification 30 

Analytical table of superfamilies 30 

Pupae with functional mandibles 34 

Micropterygoidea 35 

Eriocraniidae 35 

Pupae without functional mandibles 37 

Generalized pupae without maxillary palpi 37 

Hepialoidea 37 

Hepialidae 38 

Cossoidea 38 

Psychidae 39 

Cossidae 40 

Eucleoidea 41 

Megalopygidae 42 

Eucleidae 43 

Pyromorphidae 44 

Generalized pupae with maxillary palpi 44 

Tineoidea 44 

Prodoxidae 45 

Acrolophidae 46 

Tineidae 47 

Heliodinidae 47 

Aegerioidea 48 

Aegeriidae 49 

Tortricoidea 51 

Epiblemidae 52 

Olethreutidae 54 

Tortricidae 56 

Sparganothidae t 57 

Gracilarioidea 58 

Nepticulidae 61 

Heliozelidae 62 

Tischeriidae 63 

Bucculatrigidae 64 

Lyonetiidae 64 

Gracilariidae 65 

Phyllocnistidae 68 



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PAGE 

Specialized pupae with pilifers 69 

Pyralidoidea 69 

Pterophoridae 70 

Attevidae '^l 

Pyralididae v • ^2 

Papilionoidea ''^ 

Megathymidae 79 

Hesperiidae » 80 

Lycaenidae 83 

Papilionidae 85 

Pieridae 87 

Nymphalidae 88 

Specialized pupae without pilifers 95 

Yponomeutoidea 96 

Epermeniidae 96 

Yponomeutidae 97 

Coleophoridae 98 

Gelechioidea 98 

Lavemidae 99 

Scythrididae 100 

Gelechiidae 101 

Chrysopeleiidae 104 

Oecophoridae 104 

Stenomidae 105 

Gosmopterygidae 106 

Elaehistidae 106 

Noetuoidea 107 

Noctuidae j, . . 107 

Arctiidae 119 

Liparidae 121 

Bombycoidea 123 

Lasiocampidae 123 

Bombycidae 124 

Notodontoidea 125 

Geometridae 126 

Notodontidae 132 

Dioptidae 134 

Sphingoidea 135 

Sphingidae 135 

Saturnioidea 140 

Hemileucidae 142 

Ceratocampidae 143 

Saturniidae 144 

Phylogeny 147 

Acknowledgments 150 

Bibliography 152 

Plates 153 



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ArticItE II. — A Classification of the Lepidoptera based on Char- 
acters of the Pitpa.^ By Edna Mosher, Ph.D. 



Introduction 

It is within comparatively recent times that the immature stages of 
insects have been considered of any taxonomic value. The economic 
entomologist early realized the value of being able to recognize the 
immature stages, for in many orders of insects the larval stages alone 
were responsible for many ravages upon crops and orchards. Still the 
matter was not taken up by the systematists, and the workers in the 
field of economic entomology contented themselves by rearing the adult 
to determine the species, and then describing, perhaps all the stages, or 
more probably the larval and adult stages as being those of economic 
importance. Nowadays we are beginning to see that it is impossible 
to construct an adequate classification of •my group of insects unless 
we use every bit of information obtainable on their life history and 
habits. 

It is possible to multiply instances of the value of the larval stages 
in classification, so that one scarcely needs to cite examples; but the 
pupae have been less frequently used. There are cases, however, in 
which the only good taxonomic characters available are found in the 
pupal stage of the insect. Such instances are found among the nema- 
tocerous Diptera, particularly in the family Chironomidae. Scudder 
('89) was the first to attempt a classification of lepidopterous pupae, 
but his keys to the chrysalids were based, not on structural characters, 
but on the various projections from the body, the cuticular append- 
ages, the coloration, and the mode of suspension. 

Among the Lepidoptera a great deal of work has been done towards 
the classification of the larvae, but until 1893 nothing of importance 
had been done towards a study of the pupae. In this year Dr. T. A. 
Chapman, in a paper entitled "Some Neglected Points in the Pupae 
of Heterocerous Lepidoptera," called attention to the fact that the 
pupae possessed some remarkable taxonomic characters which might 
be used to clear up many of the disputed points in the classification 

•Contribution from the Entomological Laboratories of the University of Illi- 
nois, No. 48. 



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18 

of the order. This he endeavored to do for the groups in which 
material was available for study, and he has since published other 
articles as additional material was obtained. However, Dr. Chapman 
attempted no classification of the Lepidoptera on this basis, merely 
pointing out the pupal characters of the major groups and calling 
attention to instances in which a study of these characters would ap- 
parently alter the existing schemes of classification. 

The attention of American entomologists was called to this subject 
by Dr. A. S. Packard ('95) in a paper entitled "Attempt at a New 
Classification of the Lepidoptera." He made a new grouping of the 
order based upon pupal characters and figured a large number of 
species. His determinations of the homology of the various parts 
of the pupae studied were far from correct, and this, of course, in- 
validated many of his conclusions. 

Since that time nothing has been done in America towards a classi- 
fication of the Lepidoptera based on pupal characters. The purpose 
of the present investigation is to present such a classification as far 
as material has been available for study. There is also an attempt 
to throw some light on the relationships existing between the different 
groups. 

Changes Preceding Pupation 

The person who begins the study of pupae with the preconceived 
notion that the pupal stage is an interpolated one in the insect's life 
and that a pupa bears little or no resemblance to either larva or adult, 
will probably find abundant cause for a change of mind before his 
study is completed. In the case of Lepidoptera one is apt to think 
that no similarities could possibly exist between any of the three 
stages of the insect's development after it leaves the egg. After care- 
ful study, however, one is surprised with the resemblance between 
the stages, for it is of the highest importance in the study of any 
group to be able to homologize larval, pupal, and imaginal characters. 
This has been done to some extent in certain orders of insects, par- 
ticularly in those groups where the resemblance between the larva and 
adult is more striking than in the case of the Lepidoptera. Attempts 
have been made, however, even in this order, to homologize the 
mouth-parts of the larva and adult, and some of the larval structures 
have been homologized with certain structures in the pupa ; but appar- 
ently the idea that all three stages should be studied has been left for 
other minds to entertain. 

The first striking difference between larva and pupa is that of 
size. This difference is easily explained by the great difference in the 
size of the alimentary canal. Another striking difference is that the 



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pupa apparently lacks legs and prolegs. As will be shown later, the 
legs are always present, but folded and not in use, while the scars of 
all the prolegs remain to show their location and are very easily 
identified in the majority of cases. Many lepidopterous larvae possess 
striking tubercles and warts, and usually an abundance of setae. All 
larvae possess setae, but they are often inconspicuous. On the ex- 
posed portions of the body surface, in so far as observed, the pupa 
always retains the scars of these warts and tubercles, and the pupal 
body possesses setae arranged in most cases in the exact order in 
which they occurred in the larva. Many other structures of the larva 
can be easily identified in the pupa, and these will be discussed later. 

In the case of insects with complete metamorphosis the name pupa 
is applied to the stage of the insect in which it is more or less quies- 
cent while undergoing the changes which are necessary to fit it for 
its adult life. This word pupa, from the Latin meaning baby, was 
applied to this stage by Linnaeus from the resemblance of certain 
pupae to a baby which has been swathed or bound up, as was the 
custom in many parts of Europe at that time. This name was per- 
haps more appropriate for the pupae of the Lepidoptera than for 
those of any other order of insects because the appendages are usually 
all soldered to the thorax. 

The change from larva to pupa in the Lepidoptera has been 
observed by many workers and is full of surprises for the amateur 
who wishes to breed these insects. The caterpillar when ready to 
pupate stops feeding, and in many instances leaves the food plant and 
wanders about, often apparently in the greatest of haste. Many are 
then seen, on sidewalks, garden paths, and other traveled places, 
especially during the autumn months, when the majority of larvae are 
seeking a place to spend the winter. These larvae, if confined, will 
refuse food and many of them spin silk threads which are used to 
suspend the pupa or to form a cocoon. The alimentary canal is always 
freed of any food materials. The larval skin at this time loses its 
luster and becomes more and more wrinkled; and the body becomes 
shorter and shorter and appears swollen, which is due to the molting- 
fluid glands pouring their secretions between the outer and inner 
layers of cuticle. Some drops of a yellowish or reddish fluid are 
usually found in the place where larvae are confined and this, together 
with their peculiar appearance, often leads the amateur breeder of 
Lepidoptera to think that decomposition is taking place, and results 
in the hasty disposal of the now helpless insect. In the case of larvae 
which spin a cocoon these changes are not so easily observed, unless 
the cocoon is a very frail one, because most of the changes described 
take place inside of the cocoon. These changes may occupy but a few 



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hours, or may last for nearly a week. In the case of the common 
tomato- worm, Pro to par ce Carolina, the transformation process usually 
requires five days ; certain species of Papilio obserx'^ed took but three 
days, but the time varies much with different individuals and the con- 
ditions under which they live. 

When the molting fluid has done its work in loosening the larval 
cuticle, this splits along the meson of the thorax, and is gradually 
worked to the caudal end of the body, liberating the enclosed pupa. 
The liberated pupa is covered with a more or less transparent cuticle 
and resembles the pupa of the more generalized Neuroptera, Trichop- 
tera, and Coleoptera. In all of these orders, the insects on casting 
their larval skins show the first resemblance to the adult insect. In 
the Neuroptera, Trichoptera, and Coleoptera, the appendages, as well 
as the body, are encased in a pupal skin, are free from each other and 
the body, and together with the body segments possess considerable 
freedom of motion. This does not mean that the pupae have any 
power of locomotion ; on the contrary they are quite helpless, and for 
this reason are frequently — in common with the great majority of 
pupae — protected by some sort of a cocoon, or earthen cell. The 
lepidopterous genus Micropteryx, which is supposed by many to be 
the most generalized of its order, retains freedom of motion in all the 
appendages and in all but the fixed caudal segments of the abdomen. 
This freedom of motion is gradually lost in lepidopterous pupae as 
specialization advances, and the adult appendages are not fully de- 
veloped when the pupal stage is assumed, although the cases of the 
appendages of the pupa are fully formed. Specialization in the pupa 
consists also in the hardening of the exposed parts of the cuticle 
through the deposition of chitin, and in the soldering of the appen- 
dages to each other and to the body of the pupa. In the generalized 
families the appendages are soldered to each other but often remain 
free from the body surface; later the wings become attached to the 
body surface, but any parts of the antennae, legs, or maxillae extend- 
ing beyond their caudal margins remain free. The tips of these ap- 
pendages are provided for in various ways in the higher families, but 
are always found soldered firmly to the surface of the body of the 
pupa. Proceeding hand in hand with the soldering down of the ap- 
pendages is the loss of motion in the abdominal segments. Among 
certain families there is motion between all of the adjacent segments. 
There is, however, a successive loss of motion between segments, until 
the conjunctiva between all but two of the segments is inflexible in 
some forms, and even in some of the Lepidoptera, entire freedom of 
motion has been lost in all of the segments. 



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Among generalized forms where the appendages are soldered to- 
gether, the cuticle of the exposed parts of the body contains but very 
little chitin, and is but slightly differentiated in texture from the cuticle 
of the hidden surfaces. When the imago emerges, or even before 
that time if the body is slightly pressed, the appendages separate very 
readily from each other, and are not torn upon the emergence of the 
insect, so that the pupal skin often remains complete except for the slit 
on the dorso-meson through which the imago emerged. A very differ- 
ent condition exists, however, among highly specialized forms. Here 
the exposed portions of cuticle become very hard and firm, while those 
which are not exposed are very thin and delicate and are almost en- 
tirely destroyed at the emergence of the imago. The outer covering, 
of course, being so firmly soldered together, remains in one piece and 
is apparently complete except for the slit through which the insect 
emerged. This has led many to think that this outer chitinized por- 
tion was the entire pupal skin and that it was a structure, analogous 
perhaps to an egg shell, in which the pupa had been enclosed. 

Another remarkable difference between the generalized and the 
highly specialized Lepidoptera lies in the fact that in the latter the ap- 
pendages are not fully formed when pupation takes place, but con- 
sist of the transparent cuticular coverings through which one or more 
slender tracheae may be seen. The duration of the pupal stage doubt- 
less influences this, there being a stronger tendency among highly 
specialized forms to hibernate as pupae. 

During the life of the pupa the adult parts are developing, and be- 
fore it is time for the imago to emerge, the cuticular parts of the adult 
are fully formed. In the generalized families previously mentioned 
and in some specialized forms where the pupal cuticle remains more 
or less transparent, one is able to see a part of the development taking 
place, especially in the case of the appendages. The scales appear on 
the legs and wings and the color pattern may often be easily traced on 
the latter several days before the emergence of the insect. This stage 
of the insect, after the cuticular parts are fully formed, and while it 
still retains its pupal skin, has been designated as the preimago*. If 
the pupal skin is not already dark in color, it grows considerably 
darker in the last few days before the insect emerges, and one is thus 
able to determine when the preimago stage is reached. 

•Packard applied the term subimago to the corresponding stage in certain 
Hymenoptera. This is an unfortunate use of the term as subimago had already been 
applied to the first winged stage of the Ephemeridae. 



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22 

EXTERNAI, M0RPH0W>GY 

The most important work on pupal morphology has been done by 
E. B. Poulton and Dr. T. A. Chapman. Poulton ('91) in his paper 
on the "External Morphology of the Lepidopterous Pupa" discusses 
a few pupal structures but does not attempt to name all of the parts 
or to locate any of them. So far as known he was the first to point, 
out that the pupal structures were more than cases for the imaginal 
structures and objected to the terms pterothecae, ophthalmothecae, etc., 
as applied to pupae. Believing that Poulton's theory is correct, such 
terms have not been used in this discussion, nor the terms wing cases, 
antennal cases, leg cases, etc., but these are spoken of simply as wings, 
antennae, maxillae, etc. Chapman's papers, already referred to, dis- 
cuss very fully some of the structures and describe their exact loca- 
tion ; but as they include only a few figures one is left very much in 
doubt as to the identity of many of the structures and their location. 
W. Hatchett Jackson ('91) published a very valuable paper on the 
"Morphology of the Lepidoptera" in which he discussed the external 
determination of sex in the pupa. A short discussion of the chrysalis 
was published by Dr. S. H. Scudder ('89), and some of the parts were 
named. In a paper previously cited. Dr. A. S. Packard ('95) gives 
many figures of pupae and names the parts, but his homologies are 
far from correct. It seems necessary, therefore, before proceeding 
further, to discuss the principal pupal structures and indicate their 
location by means of figures. 

The homologies given in this paper were determined by a series 
of dissections of pupae in various stages of development, the preimago 
being found most valuable for this purpose. Pupae of nearly every 
family mentioned in this discussion have been studied in this way, be- 
ginning with the Micropterygoidea and extending through the He- 
pialoidea, Cossoidea, and other generalized families, including the 
Saturnioidea which are believed to be the most specialized of lepidop- 
terous pupae. The change from larva to pupa has been watched in 
many species and the subsequent folding and soldering down of the 
appendages carefully noted. A large number of species have been 
bred and a study of the method of dehiscence, as shown by the pupal 
skin, has thrown considerable light on many instances where there 
was doubt as to the number of free abdominal segments, or where a 
suture was obscured by folds or other modifications of the integument. 

The three regions of the body — head, thorax, and abdomen — ^are 
easily recognized, and each will be discussed in turn. There occur, 
on all of the regions of the body, in different families prominent pro- 



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jections and ridges of various types especially in the Papilionoidea. 
These projections have no morphological significance. 

THE HEAD 

The usual sclerites found in the head of generalized insects may 
be located in lepidopterous pupae. The sutures are distinct in gen- 
eralized pupae, but are obliterated in the more specialized groups. 

Vertex. — This is an area found on the dorsum of the head. It 
reaches its highest development in the Gracilarioidea, but is usually 
distinct in all generalized pupae. It is bounded cephalad hy the Y- 
shaped epicranial suture (es), and may be seen in Figures 3, 25, 29, 
33, 49, 53, and 56; V. This area was referred to by Chapman and 
Packard as the dorsal head-piece. 

front. — The front is the sclerite to which the antennae are at- 
tached. It is bounded by the epicranial suture on the dorsal surface, 
and on the ventral surface by the f ronto-clypeal suture, which nor- 
mally extends for a short distance caudad from the base of each an- 
tenna and then transversely to the median line. In some pupae where 
there is a "shoving back" of the head parts as in the Pyraustidae (Fig. 
76) and Sphingidae, the front is located on the dorsum of the head. 
The fronto-cljrpeal suture is usually not distinct except in very gen- 
eralized forms. The superfamily Gelechioidea, however, shows it 
very distinctly. It is indicated in Figures i, 8, 26, 30, and 36; f. In 
generalized pupae the front bears two setae on each side of the meson 
which are often very conspicuous. 

Genae, — These sclerites are distinctly bounded in Eriocraniidae 
and Hepialidae (Figs, i and 8, g). They are found laterad of the 
front and clypeus, and mesad of the glazed eye. The mandibles are 
always adjacent to the genae at their lateral margins. 

Clypeus, — Remarkably few pupae have the clypeus definitely 
l)Ounded. The suture between the clypeus and labrum is seldom pres- 
ent, although it is often indicated by a furrow. It is then impossible 
to determine accurately as to its presence, but it has been considered 
as if it were present. The boundaries of the clypeus are shown very 
distinctly in Figure i, cl. In the Hepialidae (Fig. 8) there is no 
clypeo-labral suture present although all the other head sutures are 
distinct. The clypeus can usually be identified by the presence of the 
invaginations for the anterior arms of the tentorium, which are asso- 
ciated with its lateral margins. This sclerite often bears prominent 
setae, and in the pupae of species whose larvae are borers it has 
often a distinct cutting plate or ridge. 



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Tentorium, — The invaginations for the anterior arms of the ten- 
torium are very distinct and are either small pores or slit-like openings. 
They are associated with the lateral margins of the clypeus and are 
distinct in most pupae (Figs, i, 14, 19, 30; at). 

Labrum. — The labrum is usually distinct along its lateral and dis- 
tal margins, but seldom separated from the clypeus by a distinct su- 
ture. Like the clypeus it usually bears setae which are especially con- 
spicuous in the Eriocraniidae (Figs, i and 2, lb). A peculiar de- 
velopment occurs in the Heliozelidae and some other families where 
the labrum extends caudad over the appendages (Fig. 50). 

Pilifers. — This term is applied to the caudo-lateral projections of 
the labrum, which are so well developed in many Lepidoptera. They 
are very large in certain superfamilies, notably the Pyralidoidea and 
the Papilionoidea, and their presence is easily detected by the lobes 
which are adjacent to the caudo-lateral angles of the labrum and often 
approximate, or meet on the meson caudad of it, or are separated by 
a narrow piece of the labial palpi (Figs. 70, 72, 74, 76, 78, 79; pf). 
The mandibles figured by Scudder ('89, Vol. 3, PI. 87, Fig. 25) are 
the pilifers. There are often well-developed pilifers present, how- 
ever, when there are no external indications of their presence. 

Mandibles. — The mandibles are always located adjacent to the 
caudo-lateral angles of the labrum. They are not functional except 
in the Micropterygoidea. In this superfamily, as shown in a pupa 
of the Eriocraniidae (Fig. i, md), the mandibles are very large and 
used by the pupa to cut its way out of its cocoon and in working its 
way to the surface of the ground. In the Hepialidae (Fig. 8, md) 
and in some other families (Fig. 11, md) the mandibular area is def- 
initely bounded. In still other families the area is distinctly elevated 
and usually rugose, as in the Eucleidae and Aegeriidae (Figs. 19 and 
36, md). This type of mandibular area is observed in many of the 
Sphingidae. In the majority of pupae, however, the mandibles are 
represented by a smooth area situated in the position indicated above. 

Eye-pieces, — These are situated laterad of the genae and mesad of 
the antennae. There are always two regions to be noted: a smooth 
mesal portion, sometimes only a narrow band but often a wider lunate 
piece, called the glazed eye-piece; and the larger lateral portion, the 
sculptured eye-piece. The latter is so called bcause it is always sculp- 
tured like the adjacent parts of the thorax. The sculpturing on the 
head is seldom like that found on the thorax and abdomen, but, 
strange to say, that on the sculptured eye-piece is always like that on 
the thorax, although the eye-piece is probably an extension of the ver- 
tex. On the dehiscence of most generalized pupae the eye-pieces are 



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separated from the face-parts and remain attached to the conjunctiva 
which joins the vertex to the prothorax (Fig. 43). In the specialized 
forms they remain attached to the face-parts. A peculiar modification 
is found in the Eucleoidea (Figs. 17, 19, 23; se, ge) in which the eye- 
pieces form movable flaps seemingly to protect and to cover the meso- 
thoracic spiracles which lie underneath. The glazed eye-piece prob- 
ably represents the pupal eye. 

Antennae. — These are always attached to the front and extend 
laterad, curving to the ventral surface of the body mesad of the meso- 
thoracic wings. They may always be identified without any trouble 
(Figs. I, 8, II, 15, 28; a). In pupae with broadly pectinate antennae, 
as the Saturniidae, the mesal portion is frequently elevated and has 
been referred to as the "stem of the flagellum" of the antennae. 

Labial Palpi. — These appendages lie adjacent on the meson caudad 
of the labrum except in the Eriocraniidae (Fig. i, Ip). They are 
visible in the majority of pupae (Figs. 8, 15, 28, 45, 61). They are 
frequently overlaid and concealed by the maxillae at their proximal 
end as in Figures 61 and 67, Ip. Often they are entirely concealed by 
the maxillae with the exception of a small V-shaped piece just caudad 
of the labrum (Fig. 72). This was thought by Scudder to be a special 
piece for covering the base of the tongue. 

Maxillae. — Where labial palpi are visible they occupy a mesal po- 
sition, caudad of the labrum, with the maxillae laterad of them. When 
they are invisible and apparently absent, the maxillae lie adjacent on 
the meson, often overlying and concealing the proximal ends of the 
labial palpi as mentioned above. The maxillae (Figs, i, 8, 17, 24, 
28; mx) vary greatly in length but are never entirely lacking or con- 
cealed in the pupa. They often extend beyond the caudal margin of 
the wings, being sometimes free and sometimes soldered to other ap- 
pendages. The greatest development is found in certain of the 
Sphingidae where the maxillae do not extend beyond the caudal mar- 
gin of the wings, but the extra length is taken up in a loop at the proxi- 
mal end which forms the so-called "jug handle'* of Sphinx pupae. The 
maxillae are always measured on the meson from the caudal margin 
of the labrum to their distal end and are usually compared in length 
with the wings, which are measured from the caudal margin of the 
labrum to their caudal margin on the meson. 

Maxillary Palpi. — Each palpus is represented on each side by a 
subrectangular or triangular area caudad of the eye-pieces and lying 
along the cephalic margins of the prothoracic and mesothoracic legs, 
frequently reaching as far mesad as the proximo-lateral angle of each 
maxilla. The normal position of these appendages is discussed under 



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the family Eriocraniidae and shown in part in Figure i, mp. They 
may also be seen in Figures 28, 30, 32, 36, 38 ; mp. Structures which 
may be maxillary palpi are found in the genus Gracilaria (Fig. 47). 
The peculiar extensions of the maxillae in the Cossoidea and Eu~ 
cleoidea are not considered as maxillary palpi (Figs. 15, 19, 23). 

THE THORAX 

The three segments of the thorax are always distinct. They are 
only visible on the dorsum, because the ventral and lateral surfaces 
are covered by the appendages. 

Pro thorax. — This segment probably varies more in size and shape 
than any of the others. There are some forms, as in the Gracilarioi- 
dea, Yponomeutoidea, and others, where the prothorax is very short 
on the meson (Figs. 53, 56, 58; p) or even invisible (Fig. 54), but is 
very wide at each lateral margin. It is longer in the Galleridae and 
certain families of Noctuoidea than in any other pupae examined. 

Pro thoracic Legs. — These lie adjacent to the maxillae at their 
proximal end. The coxae are frequently exposed, especially in gen- 
eralized pupae where the appendages are free (Figs, i, 11, 19; cxi), 
and dissection frequently showed a segment cephalad of the coxa, 
the trochantin, although there was no distinct suture indicated on the 
exterior, this being covered by the mouth-parts. The trochanter is a 
very small segment usually found at the caudal end of the femur 
when the leg is folded and is therefore generally concealed by the 
tibia and tarsus. The femur extends from the trochanter cephalad to 
the caudal margin of the head. It is frequently concealed by the tibia 
and tarsus which are the only portions of the prothoracic leg always 
visible, but they are often shoved slightly laterad so that a portion of 
the femur is exposed (Figs, i, 8, 24, 32, 36; fi). The tibia and tarsus 
are seldom divided by a suture except in generalized pupae, where all 
the segments, even of the tarsi, are readily distinguished (Fig. i, li). 

Mesothorax. — The mesothorax is usually considerably longer than 
the other segments in specialized forms, but in generalized pupae all 
the segments are more nearly ecfual. 

Mesothoracic Spiracle. — This is usually located on the dorsum be- 
tween the prothorax and mesothorax, sunk deep in the conjunctiva 
between the segments with an opening adjacent to the caudo-lateral 
angles of the prothorax. Its primitive position appears to have been 
much farther ventrad (Fig. 2, msp) and it is found in this position 
in the specialized Trichoptera. It retains this primitive position in 
the superfamily Eucleoidea and in the family Nepticulidae. The 
caudal margin possesses curious modifications in different families in 



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the way of elevated ridges, tubercles, setae, etc., and in some of the 
Papilionoidea, particularly in the families Hesperiidae and Lycaeni- 
■dae, there seems to be a definite external closing apparatus in many of 
the genera. Sometimes there is a tuft of setae ; in others, a plug or 
plate of somewhat honeycombed appearance: 

Mesoihoracic Legs, — These are folded in exactly the same man- 
ner as the prothoracic legs and the femora are very seldom exposed, 
but may be seen in Figure i, f2. The coxae are frequently visible 
(Figs. I and 48, cx2). The mesothoracic legs are usually longer than 
those of the prothorax. The tibia and tarsus of each leg are always 
exposed (Figs, i, 8, 23, 30; I2). They lie on the venter, between the 
prothoracic legs and the antennae. 

Mesothoracic Wings, — The wings of the mesothorax almost con- 
ceal those of the metathorax, except in the most generalized forms 
where the appendages are free. In most families they are the only 
wings visible on the ventral surface (Figs, i, 8, 36, 41 ; wt). 

Tegulae. — The tegulae are the large lobes which, in the adult, 
cover the proximal end of the wing. They do not form separate pupal 
pieces but are indicated in some pupae (Fig. 2, t). The tegulae are 
referred to by many authors as the patagia. The patagia are lobes of 
the pronotum which project over the mesonotum. 

Alar Furrows. — The furrows along each lateral margin of the 
mesonotum are designated as the alar furrows. They are best devel- 
oped in the Aegerioidea (Fig. 37, af ) although there are distinct de- 
pressions in many families. 

Axillary Tubercles. — In the genera Tropaea and Telea of the 
Saturniidae, there is found a large tubercle at the base of each wing, 
with sometimes an additional smaller one. The edges of these 
tubercles are strongly chitinized and somewhat roughened, and serve 
to cut the cocoon for the emergence of the moth. They are probably 
assisted in this by the peculiar development of the wing sclerites of the 
preimago, which protrude into these tubercles and are sometimes 
found to have cut the pupal skin at the apex of the tubercle. 

Metathorax. — This segment is longest in generalized forms, where 
its length is nearly equal to that of the mesothorax. 

Metathoracic Legs. — The tibiae and tarsi of the metathoracic legs 
are never normally exposed for their entire length but are concealed 
\yy the other appendages excepting at their distal end. Only a small 
portion is visible in specialized pupae, and the appendages are often 
wholly concealed (Figs. 2, 8, 36, 45; I3). 

Metathoracic Wings. — These are usually covered by the meso- 
thoracic wings except for a narrow strip along their dorsal margin. 



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In a few families a narrow strip of the metathoracic wings is visible 
on the ventral surface caudad of the mesothoracic wings (Figs, i, 8, 
19; W2). 

THE ABDOMEN 

The abdomen consists of ten segments, of which three, segments 
8-10, are always "fixed"; that is, they possess no power of indepen- 
dent motion. In the generalized forms motion is possible between all 
of the other segments. A segment is said to be movable when there is 
movement between its caudal margin and the segment caudad of it. 
In many pupae the movable segments are capable of being telescoped 
so that only their caudal margins are visible. When the cephalic 
margin of a movable segment is referred to, it includes the rounded 
part of the segment which is covered by the transverse conjunctiva 
when the segments are retracted, or telescoped. 

Proleg Scars, — The scars of the larval prolegs are found on the 
ventral surface near the meson (Fig. 11, psc) and are often conspic- 
uous. 

Tubercle Scars. — Those families in which the larvae have promi- 
nent tubercles show very definite scars in the pupae. These are espe- 
cially noticeable in the Saturniidae. 

Setae. — There are usually setae present on the abdomen, and they 
are arranged much as are those of the larvae. They are often very in- 
conspicuous, otherwise they might furnish good taxonomic characters. 
There is often a dense covering of secondary setae over the entire sur- 
face, as in some gelechiids and lasiocampids. The Pterophoridae re- 
tain a spiny armature similar to that found in the larvae. 

Spines. — These are found covering the dorsum of the abdomen 
in generalized pupae (Fig. 49), and larger ones are also found at the 
caudal end of the body (Figs. 27, 31 ). They are arranged in rows on 
the segments in Tineoidea and Tortricoidea (Figs. 27, 31, 39, 41). 

Flanged Plates. — The flanged plates are best developed in the 
pupae of borers, but are found in other pupae as well. Figure 9 shows 
them well developed on the dorsum and also shows a well-developed 
ventral plate on the seventh segment. They are usually developed 
along the cephalic margin of the segment and prevent the telescoping 
of the segments. 

Genital Openings. — In the male the genital opening is situated on 
the ventro-meson of the ninth abdominal segment. It is usually either 
a mere slit-like opening as in Figure 5, without any adjacent eleva- 
tions, or it has a distinctly elevated tubercle on each side as in Figure 
8, go, and occasionally is situated in a slight depression. In the fe- 
males there are two openings which may or may not become con- 



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fluent. These may be mere rounded pores or slit-like openings, and 
are associated apparently with the eighth and ninth segments. The 
boundary lines between segments 8 and 9, and 9 and 10 are rarely 
distinct on the meson, and where they are distinct it seems as if the 
caudal opening were associated with the tenth segment. In the more 
specialized pupae the caudal margins of the eighth and ninth segments 
are more strongly curved cephalad near the meson than in the male 
(Figs. 34 and 44, go) and the segments are dovetailed together. The 
presence of the two openings apparently represents the more general- 
ized condition (Figs. 7, 17, 28; go). They are confluent in Podosesia 
syringae (Fig. 36) and Arc hips argyrospila (Fig. 44). 

Anal Opening. — This is always situated on the meson near the 
caudal margin of the tenth segment. It sometimes shows as a cir- 
cular opening (Fig. 7, ao) but is usually slit-like (Figs. 8, 14, 17). 
It is usually surrounded on each side with several prominent wrinkles 
or folds. 

Anal Rise, — The anal opening is frequently situated on the summit 
of a mound-like elevation known as the anal rise. The setae on this 
rise are very conspicuous in certain families of Tortricoidea (Fig. 
38, ar). 

Abdominal Spiracles. — Spiracles are always present on abdominal 
segments 1-8. The spiracles of the first segment are covered, so far 
as observed, by the wings, except in the superfamily Eucleoidea and 
the family Nepticulidae. The spiracles of the eighth segment are 
never functional and show no distinct opening. 

Spiracular Furrows. — On the cephalic margin of the movable seg- 
ments cephalad of the spiracles are found furrows which frequently 
extend almost to the meson on both dorsal and ventral aspects. They 
occur in several families, as the Liparidae and Geometridae, but are 
best developed in the Sphingidae, where they are lacking in but a few 
genera. They are usually separated by sharply carinate ridges and 
are of various types, but their function is unknown. 

Cremaster. — The cremaster is a prolongation of the tenth segment 
and is not found in the more generalized pupae. It was homologized 
by C. V. Riley with the suranal plate of the larva. It is of various 
lengths and shapes and often bears setae at the distal end. Two types 
of cremaster are shown in Figures 41 and 65, cr. Its length is meas- 
ured on the ventral surface from its junction with the curve of the 
ventral surface of the body, as in Figure 44, where ab represents the 
cremastral length. 



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Classification 

As no extensive classification of the Lepidoptera based on pupal 
characters has been attempted hitherto, little has been done to deter- 
mine what characters are of value in defining superfamilies, families, 
and genera. It has been necessary, therefore, to base specific, generic, 
and other distinctions on those characters found in such material as 
could be secured. The present investigation has been limited by the 
difficulty in obtaining representatives of many groups, and it is not 
exp^ected that the tables and descriptions given will do more than 
furnish a basis for later work upon the subject. It is hoped, however, 
that they will call the attention of entomologists to the vast possibili- 
ties opened up by the use of the taxonomic characters available in 
pupae, and that further studies of the diflferent groups will make it 
possible to identify an insect in one more stage of its life cycle — which 
can not fail to be of importance in the case of our economic species. 

Analytical Table of Superfamilies 
a. Mandibles present, large, functional, decussating, and extending be- 
yond the lateral margins of the body. mICROPTERYGOIDEA. 

aa. Mandibles, if present, never large, parallel or subparallel, and 
usually represented by small elevated tubercles. 

b. Movable abdominal segments present cephalad of the fourth, or if 
no segments are movable cephalad of the fourth then the ap- 
pendages free from each other and never soldered to. the body 
wall, and the vertex longer than the prothorax measured on the 
meson. 

c. True maxillary palpi never present, but sometimes lateral exten- 
sions of the maxillae (Figs. 15 and 19). 

d. Body heavily chitinized and bearing transverse rows of spines 
or setae on the abdominal segments; spiracles never visible 
on the first abdominal segment. 

e. Mesothorax never more than twice the length of the meta- 
thorax; seventh abdominal segment with a large flanged 
plate on the ventral surface ; antennae filiform, short, only 
reaching caudad to the proximal end of the mesothoracic 
legs; head sutures all present except the clypeo-labral. 

HEPIALOIDEA 

ee. Mesothorax always more than twice the length of the 
metathorax ; seventh abdominal segment never with a large 



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flanged plate on the ventral surface ; antennae, if present, 
pectinate and reaching farther caudad than the proximal 
end of the mesothoracic legs; none of the head sutares 
distinct for the whole length. rO^^OTDFA 

dd. Body never heavily chitinized, and never bearing the spines 
or setae on the abdominal segments in rows; spiracles 
always visible on the first abdominal segment. 

EUCLEOIDEA. 

cc. True maxillary palpi usually present; if absent, then the ap- 
pendages free from each other, or the vertex longer than the 
prothorax on the meson, or the body possessing a distinct cre- 
master. 

d. Dorsum of abdomen with a covering of small spines, usually 
over the entire length of the segment and not arranged in 
distinct rows; if spines are arranged in rows then the maxil- 
lary palpi are absent; vertex always longer than the pro- 
thorax on the meson. GRACILARIOIDEA. 

dd. Dorsum of the abdomen with a distinct row of spines along 
the cephalic margin of the segment, with or without a 
caudal row ; spines seldom found elsewhere on the segment 
but, if present, then the maxillary palpi present and well 
developed. 

e. Caudal row of spines never present on the dorsum of the 
abdominal segments ; maxillary palpi always present. 

TINEOIDEA 

ee. Caudal row of spines always present on the dorsum of the 
abdominal segments ; maxillary palpi usually present. 

f. Distinct cremaster never present; setae never present on 
the anal rise; wings narrow and pointed; large spines 
always present on the venter of the tenth abdominal seg- 

^^^^' AEGERIOIDEA. 

flf. Distinct cremaster usually present, if not, then setae pres- 
ent on the anal rise ; wings broad and never pointed ; 
large spines never present on the venter of the tenth 
abdominal segment. TORTRICOIDEA. 

bb. Movable abdominal segments never present cephalad of the 
fourth; appendages never free from each other and usually 
soldered to the body wall. 



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c. Lobes indicating the presence of pilifers always present except 
in Gallerinae (Fig. 69) and Oeneinae (Fig. 80). 

d. Maxillary palpi usually present, if absent, then abdominal 
segment seven is movable in the male, the body covered with 
a spiny armature, and both prothoracic and mesothoracic legs 
extending cephalad between the eye-pieces and the antennae, 
the former reaching nearly to the cephalic margin of the 
glazed eye, or a deep furrow lined with setae present on the 
dorsum between the ninth and tenth abdominal segments; 
antennae never clubbed at the distal end ; femora of the pro- 
thoracic legs usually visible ; labial palpi very seldom visible 
except as a small triangular or polygonal area caudad of 
the labrum and between the pilifers. pyrat TDOTDF A 

dd. Maxillary palpi never present; antennae always clubbed at 
the distal end; femora of the prothoracic legs never visible; 
a deep furrow lined with setae never present on the dorsum 
between the ninth and tenth abdominal segments; labial 
palpi never visible except as small triangular or polygonal 
areas caudad of the labrum between the pilifers and often 
entirely concealed. PAPILIONOIDEA. 

cc. Lobes indicating the presence of pilifers never present. 

d. Mesothoracic wings on the ventral surface at meson usually 
extending considerably beyond the caudal margin of the 
fourth abdominal segment, if not, then the body depressed, 
mostly in the thoracic region, the incisions between the mov- 
able segments very deep on the dorsum and venter and less 
deep at the lateral margins, and the caudal part of the an- 
tennae always adjacent on the meson for a considerable dis- 
tance ; abdominal segments 1-4 usually longer than the other 
segments ; epicranial suture always present. 

e. Maxillary and labial palpi present and well developed and 
a large portion of the prothoracic femora always exposed ; 
if maxillary palpi are not present th^i the f ronto-cljrpeal 
suture never i-isible; prothorax distinctly shorter on the 
meson than at each side, so that each half is triangular in 
outline ; appendages soldered to each other but not to the 
body wall; fronto-clypeal suture never visible; antennae 
with the caudal portion very rarely touching and not 
moniliform in appearance. yPONOMBUTOIDEA. 

ee. Maxillary palpi usually present, but labial palpi and pro- 
thoracic femora seldom visible, if visible, then the f ronto- 
cljrpeal suture distinct ; prothorax usually the same length 



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on the meson as at each side so that each half is subquad- 
rangular in outline ; appendages usually soldered firmly to 
each other and to the body wall ; body usually ovate in out- 
line, broadest in the thoracic region and usually strongly 
depres^d ; f ronto-clypeal suture usually visible ; antennae 
usually moniliform in appearance, the caudal portion al- 
ways adjacent on the meson, usually for some distance ; if 
only touching, then the f ronto-clypeal suture is distinct. 

GELECHIOIDEA. 

dd. Mesothoracic wings on the ventral surface of the body at. 
meson rarely extending beyond the caudal margin of the 
fourth abdominal segment, if beyond, then maxillary palpi 
never present ; abdominal segments 1-4 or 1-6 rarely longer 
than the other segments; epicranial suture seldom visible. 

e. Labial palpi usually present and well developed and from 
one fourth to one fifth the length of the wings, if not vis- 
ible then the body usually shaped as in Figure 104; the 
abdomen with setae arranged around the scars of larval 
verrucae, and usually flanged plates on the abdomen and a 
cremaster present ; or with a more or less dense covering of 
setae never arranged around larval verrucae, the body 
never of the shape in Figure 104 and flanged plates never 
present on the abdomen nor a distinct cremaster. 

f . Labial palpi usually present and well developed and the 
prothoracic femora usually exposed, or if not, then both 
prothoracic and mesothoracic legs reaching cephalad to 
the eye-pieces ; if both labial palpi and prothoracic fem- 
ora are wanting then the body of the type in Figure 
104 ; the abdomen with setae arranged around the scars 
of larval verrucae, flanged plates usually present on the 
abdominal segments and a distinct cremaster often 
present; body never with a more or less dense cover- 
ing of setae, except arranged as mentioned above ; max- 
illary palpi occasionally present. NOCTUOIDEA 

ff. Labial palpi sometimes present, body never with a cre- 
master and always with a more or less dense covering 
of setae which are never arranged around larval ver- 
rucae; prothoracic femora never exposed; maxillary 
palpi never present. BOMBYCOIDEA. 

ee. Labial palpi never visible, unless represented by small tri- 
angular or polygonal areas caudad of the labrum; body 
very seldom with visible setae. 



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f. Suture adjacent to the proximal ends of the antennae 
and separating the elypeus and front always present 
and very distinct; antennae never broadly pectinate 
so that the width is one fifth of the length; spiracular 
furrows often present. 

g. Antennae usually considerably broader near the prox- 
imal end, their greatest width usually greater than 
that of the prothoracic legs ; antennae usually more 
than three fourths the length of the wings, if not, then 
the epicranial suture is present, or the cremaster is 
wanting, or if present, bifurcate at the distal end or 
bearing hooked setae ; dorsum of the abdomen usually 
with a deep furrow between the ninth and tenth seg- 
ments; scar of a caudal horn never present on the 
dorsum of the eighth abdominal segment ; labial pal- 
pi sometimes visible as small triangular or polygonal 
areas caudad of the labrum. nOTODONTOIDEA. 

gg. Antennae rarely very much broader near the proxi- 
mal end, usually filiform, their greatest width seldom 
greater than that of the prothoracic legs, if greater 
then the cremaster is never wanting, nor bifurcate, 
nor with hooked setae; antennae never more than 
three fourths the length of the wings; epicranial 
suture never present ; dorsum of the abdomen never 
with a deep furrow between the ninth and tenth seg- 
ments ; scar of a caudal horn usually present on the 
dorsum of the eighth abdominal segment ; labial pal- 
pi never visible. SPHINGQIDEA. 

ff. Suture adjacent to the proximal ends of the antennae 
and separating the front and elypeus obsolete for the 
greater part of its length; antennae always broadly 
pectinate and the width at least one fifth of the length 
and often wider; spiracular furrows seldom present. 

SATUBNIOIDBA. 

PUPAE WITH FUNCTIONAL MANDIBLES 

Among the Trichoptera, from which the Lepidoptera are supposed 
to have descended and to which they are known to be very closely 
related, there are many pupae which have functional mandibles. 
They function, though, merely to assist the pupa to escape from the 
cocoon. Among the generalized Lepidoptera the pupae of one super- 



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family, the Micropterygoidea, have large mandibles which sei-ve the 
same purpose as in the Trichoptera. 

SuPEBPAMiLY MICROPTERYGOIDEA 

The most generalized lepidopterous pupae known belong to the 
superfamily Micropterygoidea, which includes two families, the 
Micropterygidae and the Eriocraniidae, characterized by the presence 
of functional mandibles. Except for a description of the fragments 
of the head by Chapman, no pupa of the Micropterygidae has been 
described, but this family is undoubtedly the most generalized, because 
the adults possess functional mandibles. 

The first complete life history of any American species of Erio- 
craniidae was worked out by Busck and Boeving and published in the 
Proceedings of the Entomological Society of Washington in 19 14 
(Vol. XVI, pp. 15 1 -163). These authors gave a short description of 
the pupa and included some excellent figures. A more detailed de- 
scription is given here as this species furnishes a working basis for 
the study of all other lepidopterous pupae. This was made possible 
by the generosity of Dr. L. O. Howard, Honorary Curator of Insects, 
U. S. National Museum, who donated some excellent material of 
Mnemonica auricyanea collected this year by Mr. August Busck at 
Falls Church, Va. 

The pupae of this species (Figs, i, 2, 3) are very small, averaging 
3 mm. in length in the males and 4 mm. in the females. The body is 
covered by a thin transparent cuticle, which shows all the imaginal 
parts in mature pupae, ma'king it exceedingly difficult to distinguish 
pupal structures from similar structures in the adult. It is also very 
difficult to determine the number and position of the setae. 

The head shows all the sutures usually present in generalized in- 
sects. The vertex is short, the epicranial suture fairly distinct and 
extending to the lateral margins of the head. The f ronto-clypeal 
suture extends transversely between the caudo-lateral angles of the an- 
tennae. The front bears two long straight setae on each side of the 
meson about half-way between the antennae and the cephalic margin 
of the head. In the middle of the cephalic aspect, between the an- 
tennae, arises a long, fleshy, beak-like projection which contains the 
long tuft of hairs present in the adult. Just caudad of the front is 
the clypeus and laterad of these are the genae in the usual position 
for the Lepidoptera. The suture between the clypeus and labrum is 
broad and somewhat chitinized, and closely appressed to its ental sur- 
face is the tentorium, to which the mandibles are attached. The 



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labrum is a large fleshy projection bearing on its ectal surface six pairs 
of very long setae which extend beyond the lateral margin of the body, 
and on the ental surface two groups of much shorter setae, which 
project slightly beyond its caudal margin. All of the appendages of 
the head are free. The labial palpi are rather short, with three seg- 
ments, and are somewhat enlarged and blunt at the distal end. The 
mandibles are exceedingly large and are attached to the ental surface 
of the clypeus, extending beyond the lateral margin of the body. They 
are heavily chitinized and serrate along the cephalo-lateral margin. 
The distal end is broadened and thickened, somewhat circular in out- 
line, concave and strongly toothed. The maxillae are short and the 
halves are widely separated. Each half is strongly bent near the dis- 
tal end, which is directed cephalad and mesad. The maxillary palpi 
are long, apparently with six segments, and pass from the mouth 
dorsad and then out towards the lateral margin of the head, making 
a series of curves which finally bring them between the eyes and the 
antennae. The distal end is folded close to the body and lies just 
caudad of the eye. The antennae show a long pedicel with many 
shorter segments and extend for more than half the length of the 
wings. 

The thoracic segments are all more or less movable. The thorax 
is short, strongly elevated, and moves freely, the greater part of its 
exposed portion being conjunctiva. The mesothorax and metathorax 
are nearly equal in length, but seem to possess little power of inde- 
pendent motion. On the dorso-meson of these two thoracic segments 
and the first abdominal segment is found a strap-like cuticular thick- 
ening which is apparently for strengthening the thorax. The tegulae 
are indicated by the dotted lines in Figure 2 because they do not seem 
to be distinct pupal structures. The thoracic appendages are also free. 
All of the coxae are visible and usually the femora of the prothoracic 
and mesothoracic legs. The metathoracic legs are usually hidden be- 
neath the wings except at the distal end, which normally curves around 
the caudal end of the body. The wings never extend to the caudal 
margin of the body. 

The first seven abdominal segments are movable in both sexes. 
The remaining segments are not distinctly sutured and possess no 
power of independent movement. The genital openings are rather 
difficult to locate. That of the male is found as a slit-like opening 
on the ventro-meson of the ninth segment (Fig. 4). There are two 
openings in the female (Fig. 7, go), apparently located on the ventro- 
meson of the eighth and ninth abdominal segments. 



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The tenth segment is longer in the female than in the male, pre- 
sumably on account of the ovipositor. The females always have the 
eighth, ninth, and tenth segments curved ventrad and closely ap- 
pressed to the ventral surface of the body. This is shown where the 
caudal segments are slightly separated from the body, in Figure 2. 
Figures 6 and 7 give dorsal and ventral views of these caudal seg- 
ments, and Figures 4 and 5 show the same segments of the male. The 
anal opening in both sexes is found near the caudal end of the body 
on the tenth segment. The spiracles are small, circular, and not pro- 
duced. The mesothoracic spiracle is situated in the conjunctiva con- 
necting the prothorax and mesothorax. Functional abdominal spira- 
cles are visible on segments 2-7. The dorsum of the abdomen is 
practically covered by very minute spines arranged in groups. 

The following species was examined : 
Mnemomca auricyanea Walsingham. 

PXTPAE WITHOUT FUNCTIONAL MANDIBLES 

This group includes all the superfamilies of Lepidoptera known, 
except the Micropterygoidea. In many of the other families the 
pupae possess mandibles, but they are functionless, and only indi- 
cated as small parallel tubercles or lobes. 

Oeneralized pupae without maxillary palpi 

The Hepialoidea, together with the Cossoidea and Eucleoidea, 
differ from all other generalized pupae possessing free abdominal seg- 
ments cephalad of the fourth, because of the absence of the maxillary 
palpi. Some of the families included here possess lateral prolonga- 
tions of the maxillae which resemble maxillary palpi (Figs. 15 and 19) 
and have been considered as such by some authors. These prolonga- 
tions never separate from the maxillae at dehiscence, and dissection 
has failed to find any maxillary palpi present in the mature pupae. 
None of these superfamilies possess all of the sutures found in the 
head of the generalized type, and none of them show the long, seg- 
mented antennae present in the Eriocraniidae. 

SUPERFAMILY HEPIALOIDEA 

This includes a single family, Hepialidae, of which the known 
larvae are borers. The species in this country are of rare occurrence. 
Their larvae are borers in the stems of shrubs or trees. In Europe 
some of the species are abundant and injurious. The specimens of 



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Sthenopis thule were obtained through the courtesy of Mr. J. M. 
Swaine, of the Canadian Department of Agriculture, who obtained 
them from the stems of willow at MacDonald College, Quebec. 

Family Hepialidae 

The pupae of this family are very generalized as to the number of 
sutures present in the head, the number and arrangement of append- 
ages, the comparative length of the mesothorax and metathorax, and 
the nearly equal length of the first seven abdominal segments. These 
characters are easily seen in Figures 8-10 and need no further de- 
scription. The pupae are, however, exceedingly specialized as to the 
chitinization of the body, the spines, toothed ridges and cutting plates 
on the abdominal segments, and, more than all, in the soldering down 
of all the appendages to each other and to the body, exactly as in the 
most specialized of pupae. The head, thoracic segments, and the first 
two abdominal segments are firmly soldered together, but abdominal 
segments 2-7 are free in the male and 2-6 in the female. 

The only consolidation of the head parts is that of the clypeus 
and labrum, between which the suture has been lost. The antennae, 
as well as all the other appendages, are very short in comparison with 
the length of the body. These pupae are of considerable size, that 
of Sthenopis thule being about 30 mm. in length. The larvae as far 
as known are borers, and their pupae have special adaptations for cut- 
ting their way to the surface. The most peculiar of these adaptations 
is the ventral plate on the seventh abdominal segment (best seen in 
Figure 9), which has not been found in any other pupae examined. 
The sharp ventral projections on the front also serve as cutting sur- 
faces, but similar projections are found in many pupae, particularly 
among other species whose larvae are borers and in very many of the 
leaf-mining species. The opening of the mesothoracic spiracle has 
reached the normal position for most lepidopterous pupae, being be- 
tween the prothorax and the mesothorax at each caudo-lateral angle 
of the former. The genital opening is found in the male on the meson 
of the ninth segment between two slightly elevated tubercles. In the 
female there is a single opening apparently on the eighth segment. 

The following species was examined : 
Sthenopis thule Strecker. 

SUPERFAMILY COSSOIDEA 

The pupae of this superfamily are less generalized as to head 
parts than the Hepialoidea, but nevertheless resemble them very 
closely in size, shape and arrangement of the appendages, in the num- 



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39 

ber of free segments, and in the fact that all the appendages are 
firmly soldered to each other and to the body. The antennae, how- 
ever, are of a different type, being pectinate in the Cossoidea. The 
metathorax varies considerably from the generalized t)rpe, being very 
much shorter, so that the mesothorax is about four times its length, 
measured on the median line. Many of the species in this superfamily 
have larvae which are borers, and many of the pupae are fitted to work 
their way to the surface of a burrow. However, the ventral plate of 
the seventh abdominal segment, which is so distinct in the Hepialoi- 
dea, is not present. All of the pupae have some of the body segments 
armed with spines and strongly toothed chitinized ridges, and a 
strong ridge or projection is generally present on the head. The 
families may be separated as follows : 

a. Abdominal segments 2-6 movable in the female and 2-7 in the male ; 
dorsum of abdominal segments armed with a row of sharp spines 
on the cephalic margin, and a row of setae, which are directed 
cephalad, on the transverse conjunctiva at the caudal margin; 
females without wings and antennae and larva-like in appearance. 

PSYCHIDAE. 

aa. Abdominal segments 3-6 movable in the female and 3-7 in the male ; 
dorsum of abdominal segments armed with a toothed ridge along 
each margin ; sexes similar CossroAE. 

Family Psychidae 

In this family there are no sutures apparent on the head except 
between the clypeus and labrum and the mandibles. The antennae 
are short and pectinate. The prothorax is longer and the meta- 
thorax much shorter than in the Hepialoidea, to which, however, 
these pupae show many resemblances. The dorsum of the abdomen 
has toothed chitinized ridges along the cephalic margin of some of 
the segments and rows of setae along the caudal margin on the trans- 
verse conjunctiva. The caudal end of the body bears two large, 
strong hooks directed ventrad. This description applies mostly to 
the males (Figs. 11-13), as the females are quite different as seen in 
Figure 14. 

The females never leave the cocoon during their entire life and 
have no provision for locomotion, even in the adult. It is an aston- 
ishing fact that no pupal wings are developed, because in all other 
families where the adult females are apterous the pupal wings are 
always developed, sometimes as much as in the males. Neither are 
there any pupal antennae present, no eye-pieces, nor traces of maxillae. 
The labrum and mandibles show very distinctly, both being consid- 



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erably elevated. The legs are scarcely developed, 'being represented 
by transverse chitinized elevations on the venter of the thoracic seg- 
ments. The abdominal segments are much as in the male. They 
show on the venter the proleg scars, on the dorsum the rows of 
toothed chitinized ridges and setae, but the body setae are much 
smaller and difficult to distinguish and are not represented in the 
figure. A single genital opening is found in the female, on the eighth 
abdominal segment. No hooks are present at the caudal end of the 
body. The abdominal spiracles are present on the first eight abdom- 
inal segments, but there is no visible opening for the mesothoracic 
spiracle in either sex. The only genera available for study were 
Thyridopteryx and Oiketicus. These resemble each other very closely 
and the difference between the pupae can hardly be considered as 
generic. The pupae of Oiketicus are larger and stouter, the males 
examined averaging i8 mm. in length, while those of Thyridopteryx 
were slenderer and only 15 mm. in length. The two genera may be 
separated thus : 

a. Abdominal segments 2-6 with a caudal row of setae, the row on the 
the sixth interrupted and shorter than the other rows ; caudal spines 
stout and simple; spiracles scarcely produced beyond the surface 
of the body except at their cephalic margins. 

Thyridopteryx Stephens, 
aa. Abdominal segments 2-5 with a caudal row of setae, no row on the 
sixth ever present ; caudal spines slender and with a distinct tooth ; 
spiracles distinctly produced beyond the surface of the body. 

Oiketicus Guilding. 
The following species were examined : 
Thyridopteryx ephemeraeformis Haworth 
Oiketicus abbotii Grote 

Family Cossidae 

The Cossidae are borers in the larval stage and seem to be very 
closely related to the Hepialidae, although they resemble them less 
than do the Psychidae. This family has segments 3-7 of the abdomen 
free in the male and 3-6 in the female. There is another sexual dif- 
ference to be noted, viz., the presence of an extra row of spines on 
the abdomen of the male. In this sex the seventh segment has two 
rows of spines and the succeeding segments one row; in the female 
the sixth is the last segment with two rows, the remaining caudal seg- 
ments having but one row. The epicranial suture is not distinct in 
any species, but at dehiscence Prionoxystus robiniae shows a small 
piece of the vertex on each side of the meson, and this with the con- 
junctiva bears the eye-pieces. The lateral part of the fronto-clypeal 
suture is distinct and the clypeo-labral suture is always visible. 



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This family is usually divided into subfamilies of which two, the 
Cossinae and Zeuzerinae, are discussed here. Figure 15 shows the 
ventral surface of the head and its appendages in a member of the 
Cossinae, the arrangement of the other parts being the same as in 
the Zeuzerinae (Fig. 16). 

The maxillae have prominent lateral projections in Cossinae which 
resemble maxillary palpi. These always adhere to it at dehiscence 
and are not found in Zeuzerinae. Only one genus of each subfamily 
was studied. The pupae are very large, those of Prionoxystus robin- 
iae and Zeiizera pyrina being respectively 45-50 mm. and 30-35 mm. 
in length. The two genera studied may be separated as follows : 

a. Head without a prominent cephalic projection; maxillae with an 
apparently segmented lateral projection on each side resembling a 
maxillary palpus, but adhering to the maxillae at dehiscence; an- 
tennae more than half the length of the wings and gradually taper- 
ing ; abdominal segments with the cephalic ridges much larger than 
the caudal ones and armed with long even teeth. 

Prionoxystus Grote. 

aa. Head with a prominent cephalic projection, maxillae never with an 
apparently segmented lateral projection on each side ; antennae less 
than half the length of the wings and narrowed abruptly near the 
middle J abdominal segments with the cephalic and caudal ridges 
similar, the teeth short and uneven Zeuzera LatreiUe. 

The following species were examined : 
Prionoxystus robiniae Peck 
Zeuzera pyrina Linnaeus 

SUPERFAMILY EUCLEOIDEA 

The pupae of this superfamily are quite specialized as to the head 
parts, the epicranial suture being the only one visible in all the 
families. They have followed a very different line of development 
from the Cossoidea and Hepialoidea, because all of the generalized 
families retain freedom of motion between all the segments except 
those fixed at the caudal end of the abdomen, and between all of the 
appendages. The cuticle is very thin and transparent in almost all 
genera and the dorsum of the abdominal segments in all of them has 
a covering of small spines over the greater part of the segment. All 
of the families show the spiracles distinctly on the first abdominal 
segment. The only other family in which this was observed, the 
Nepticulidac, has a well-developed maxillary palpus. The meso- 
thoracic spiracle of each side is in a rather unusual position in this 



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superfamily. The opening is on the dorsum in the normal position, 
and is very large, with a strongly arched cephalic margin; but the 
spiracle is on the ventral surface directly under the sculptured eye- 
piece in Megalopygidae and Eucleidae, and a little farther laterad in 
Pyromorphidae so that it comes partly under the antennae. The 
spiracle, with the adjoining parts slightly pushed aside to show their 
relation, is seen in Figure 21. The family Pyromorphidae being more 
specialized than the other two families differs from them considerably, 
but its relationship to them is evident. The three families included 
here may be separated as follows : 

a. Dorsum of abdominal segments with spines on the cephalic part and 
a covering of coarse setae on each caudo-lateral part which does not 
usually extend to the meson; maxillae simple quadrangular pieces, 
without any lateral prolongations; a large conical tubercle caudad 
of each abdominal spiracle on segments 2-6 ; mesothorax never ex- 
tending caudad to the first abdominal segment Meoaloptoidae. 

aa. Dorsum of abdominal segments with short spines, but never with a 

covering of coarse setae on any part; tubercles never present caudad 

of any of the abdominal spiracles. 

b. Labial palpi present; mesothorax with a long mesal lobe reaching 

on to the first abdominal segment ; maxillae always less than half 

the length of the wings Pyromorphtoae. 

bb. Labial palpi absent; mesothorax never with a long mesal lobe 
reaching on to the first abdominal segment ; maxillae more than 
half the length of the wings Pyromorphtoae. 

Family MEOALOPYoroAE 

The Megalopygidae have the head and thoracic segments free, also 
abdominal segments 1-7 in the male and 1-6 in the female. The ap- 
pendages are entirely free from each other and from the body wall. 
The body is soft and covered with a thin, delicate, transparent cuticle 
which is slightly chitinized. There are always setae on the dorsum 
of the abdominal segments as well as spines. The setae are found on 
each side of the meson on the caudal half of all the segments. The epi- 
cranial suture is distinct but all the other head sutures are obliterated. 
The front has a distinct projection and the mandibles show as dis- 
tinctly elevated tubercles. The size and arrangement of the parts 
may be seen in Figures 17 and 18. This family together with the 
Eucleidae possesses a very peculiar eye-piece. Chapman ('94, p. 349) 
called attention to this structure and spoke of it as the "eye-flange". 
This eye-piece, in reality the sculptured portion, is free along its 
lateral and caudal margins and extends well out on to the antennae. 



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43 

It is, however, much more wrinkled and sculptured than any other 
portion of the body. These eve-pieces move up and down in living 
pupae during respiration and allow one to see the mesothoracic 
spiracle omdemeath. The mesothorax possesses some well-defined 
alar ridges and its caudal margin extends in a broad curve nearly to 
the caudal margin of the metathorax. The large conical tubercles are 
found caudad of the spiracles on abdominal segments 2-6. The body 
of Lagoa crispata Packard, the only species studied, is strongly arched 
on the dorsum of the abdomen and is short and thick-set. Its length is 
about 18 mm. and the greatest breadth 10 mm. 

The following species was examined : 
Lagoa crispata Packard. 

Family Eucleidae 

The Eucleidae retain the same movable segments as the family 
Megalopygidae, which they strongly resemble. The pupae of Eucle- 
idae, however, are usually only half the size of the latter, averaging 
about 9 mm. in length. They also retain the same head sutures, but, 
as in Prolimacodes, they often show a distinct furrow marking the 
position of the lateral part of the fronto-cl)rpeal suture. The eye- 
pieces are identical with those described for Megalopygidae. The 
size and arrangement of parts may be seen in Figures 19, 20, and 23. 
In two of the genera studied, Sibine and Euclea, the maxillae, in ad- 
dition to the usual cephalo-lateral extension found throughout the 
family (Fig. 23), have peculiar modifications in the form of long 
lateral prolongations extending to the antennae. Usually only the dis- 
tal end of this prolongation is seen between the eye-piece and the an- 
tennae, as in Figure 19, the dotted line showing the connecting part. 
These two genera also have a distinct groove in each half of the max- 
illae, into the caudal part of which the femur of the prothoracic leg is 
fitted. The cephalic margin of the pronotum has a distinct median 
notch, which makes it appear bilobed, and each lobe is prolonged 
ccphalad over the caudal margin of the head (Fig. 22). The meso- 
notum is prolonged into a rounded or pointed lobe which reaches on 
to the first abdominal segment. Only three genera were available for 
study. These may be separated by the following table : 

a. Maxillae never with a lateral projection reaching to the antennae; 
mesothorax with a strongly carinate median line; caudal lobe of 

the mesonotum broadly rounded Prolimacodes Schaus. 

aa. Maxillae with lateral projections reaching to the antennae ; meso- 
thorax never with a strongly carinate median line. 

b. Mesonotum with the caudal lobe pointed Euclea Hiibner. 

bb. Mesonotum with the caudal lobe broadly rounded, almost trun- 
cate Sibine Herrich-Schaeflfer. 



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The following species were examined : 
Prolimacodes scapha Harris 

Bticlea dclphinii Boisduval, chloris Herrich-Schaeffer 
Sibine stimulea Clemens 

Family Ptromorphidae 

This family is much more specialized than cither the Megalopy- 
gidae or the Eucleidae and resembles them but little. The body is 
flattened and has lost the power of motion except in the abdomen. 
Abdominal segments 2-7 are free in the male and 2-6 in the female. 
The appendages are also very slightly soldered together. The pres- 
ence of spines on the abdominal segment, together with the absence 
of maxillary palpi, is considered sufficient evidence that it belongs to 
the superfamily Eucleoidea. Figures 24 and 25 show the essential 
points of its structure. The only genus available for study was 
Harrisina. 

The following species was examined : 
iHarrisina americana Guerin-Meneville. 

Cteneralized pupae with maxillary palpi 

The remaining pupae which retain either free segments cephalad 
of the fourth abdominal segment or free appendages, have followed 
two distinct lines of development. In the first group the generalized 
condition of the body found in the Eriocraniidae has been retained as 
to comparative length of segments and the covering of the dorsum of 
the abdomen with fine spines. The metathorax is nearly always more 
than half the length of the mesothorax, while the prothorax tends to 
become shorter at the meson and broader at the lateral margins, so 
that each half appears triangular. In the second group, the covering 
of spines on the dorsum of the abdomen has been gradually changed 
and there is one very well-developed row of spines at the cephalic mar- 
gin of each segment, with or without a similar caudal row. In this 
group the prothorax is longer and somewhat quadrangular in shape 
and the metathorax is relatively shorter. This group includes the 
superfamilies Tineoidea and Tortricoidea, and being much smaller 
than the other will be considered first. 

Superfamily TINEOIDEA 

The families included here possess one row of spines along the 
cephalic margin of the dorsum of the abdominal segments, and well- 
developed maxillary palpi. In one family, Prodoxidae, the primitive 



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covering of fine spines has been retained, but it may be easily differ- 
entiated from all other pupae bearing spines of two sizes in a similar 
position on account of the large maxillary palpi. 

The family Heliodinidae is included here for the sake of con- 
venience as it possesses only the cephalic row of spines on the dorsum 
of the abdominal segments. It is, however, much more nearly related 
to the Tortricoidea. The families Prodoxidae and Acrolophidae are 
more nearly related to the Tineidae. Of these the Prodoxidae are 
undoubtedly the most generalized, retaining more head sutures and a 
greater number of free segments, in addition to the spines mentioned 
above. The Acrolophidae are more generalized than the Tineidae in 
the matter of free segments, but have the appendages firmly soldered 
to each other and to the body wall. This is probably due to the fact 
that thi? larvae are sod-borers and that the pupa works its way to the 
surface. The families may be separated as follows: 

a. Mesonotum not produced into a long caudal lobe ; mesothorax seldom 
more than twice the mesal length of the metathorax. 
b. Abdominal segments 2-7 movable ; dorsum of abdominal segments 
with a covering of spines on the caudal part ; maxillae more than 

twice as long as the labial palpi Prodoxtoae. 

bb. Abdominal segments 3-7 movable ; dorsum of abdominal segments 

never with a covering of spines on the caudal part; maxillae 

shorter than the labial palpi. 

c. Antennae never extending to the caudal margin of the wings ; 

wings broadly rounded; appendages firmly soldered to each 

other and to the body ; a lateral projection never present on 

each side of the tenth abdominal segment Acrolophtoae. 

cc. Antennae extending beyond the caudal margin of the wings ; 
wings pointed; appendages only slightly soldered together 
and separating at the slightest touch; tenth abdominsJ seg- 
ment with a prominent lateral projection on each side end- 
ing in a spine Tinetoae. 

aa. Mesonotum produced into a long caudal lobe ; metathorax never 
more than one fourth the mesal length of the mesothorax. 

Heuoddodae. 
Family Prodoxtoae 

In this family abdominal segments 2-7 are free in both sexes. The 
head shows the epicranial suture plainly, and dehiscence always takes 
place on the front of the head along what is apparently the fronto- 
cl)rpeal suture, at least for a part of the distance, as shown in Figure 
26. The front bears a prominent chitinized projection armed with 
two stout teeth. The lateral margin of the eye-piece extends on to 
the antenna for a very short distance. The appendages are very 



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slightly soldered to each other, but scarcely to the body wall, and sep- 
arate very easily. The lateral view, Figure 27, shows the relative 
length of the segments. The abdominal segments, although they have 
developed a prominent cephalic row of spines on the dorsum, still re- 
tain the covering of very fine spines on the remainder of the seg- 
ment. The eighth abdominal segment bears a pair of very stout hooks 
at the apices of rounded tubercles (Fig. 27a). The pupae examined 
measured about 10 mm. in length. 

The following species was examined: 
Prodoxus quinquepunctella Chambers. 

Family Acrolophidae 

In this family segments 3-7 of the abdomen are movable in both 
sexes, but the appendages are quite firmly soldered to each other and 
to the body wall so that they do not readily separate even at dehiscence. 
There is probably also dorsal movement of the second segment, as the 
conjunctiva is well developed and both the first and second segments 
separate at dehiscence. The larvae of members of this family are sod- 
borers and it seems quite natural that pupae with this mode of life 
should have their appendages soldered down at a much earlier stage 
than those of the leaf-miners, for instance, or of the pupae that live in 
cocoons. There are none of the small spines of the generalized type 
present on the dorsum of the abdomen in this family, but a well- 
developed row of spines at the cephalic margin of the segments. There 
are also short lateral and dorsal projections of the tenth segment, with 
very sharp chitinized edges, which are evidently to aid the pupa in 
working its way to the surface. The head bears a strongly chitinized 
transverse ridge near the cephalic margin of the ventral surface. 
Figures 28 and 29 show the arrangement of parts in a pupa of this 
family in which there is a remarkable development of the labial palpi. 
The pupae are from 15-20 mm. in length. The genera may be sepa- 
rated as follows : 

a. Labial palpi never with distinct cutting plates near their proximal 
margin, the palpi not extending much over half the distance to the 
distal ends of the prothoracic legs ; two pairs of coxae visible ; spines 
of the abdominal segments long and narrow. 

Hypocolpus Walsingham. 

aa. Labial palpi with distinct cutting plates near their proximal margin, 

the palpi extending as far as the distal ends of the prothoracic legs ; 

a single pair of coxae visible ; spines of the abdominal segments 

triangular Pseiidanaphora Walsingham. 



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The following species were examined : 
Hypocolpus mortipennellus Grote 
Pseudanaphora arcanella Clemens 

Family Tineidae 

In this family the free abdominal segments are 3-7 in the male; 
no females were available for examination. Segments 1-3 separate dor- 
sally at dehiscence. The appendages are very slightly soldered to- 
gether and all separate readily except the metathoracic legs and an- 
tennae, which extend beyond the caudal margin of the wings and are 
quite firmly fastened together, the legs being underneath the antennae. 

The appendages are also slightly soldered to the body as far as the 
third abdominal segment. The arrangement of parts is shown in 
Figures 30 and 31. The fronto-clypeal suture is indicated by a clear 
line in the otherwise fairly well-chitinized cuticle. Segments 3-8 of 
the abdomen bear a cephalic row of spines on the dorsum directed 
caudad, while the ninth segment bears an interrupted group of spines 
directed cephalad. There are none of the fine spines of the generalized 
type of pupa present in this family. The tenth abdominal segment 
shows a prominent lateral projection on each side, ending in a spine. 
The setae of the body are very conspicuous. The pupae are about 4 
mm. in length. 

The following species was examined : 
Tinea pellionella Linnaeus. 

Family Heliodinidae 

This family has usually been associated with the Yponomeutidae, 
but it seems from pupal characters to be more closely related to the 
tortricids. It is very similiar to these in arrangement of parts; the 
Heliodinidae, however, have longer maxillae and they plainly show 
that dorsal motion is possible between the second and third abdominal 
segments. There are also curved setae at the caudal end of the body 
in the genus Brenthia (Figs. 32 and 33) strongly resembling those 
found in the Epiblemidae. Choreutis (Figs. 34 and 35) has a small 
dorsal plate on the tenth segment with a strong seta at each end which 
appears to represent an early state in the development of a cremaster. 
The possession of a single row of dorsal spines on the abdominal seg- 
ments, however, is like the remainder of the Tineoidea, and it is easier 
to classify them as such. They differ from the remainder of the super- 
family in having one more free segment in the male, abdominal seg- 
ments 3-7 being free in the male and 3-6 in the female. The thorax 



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differs markedly, too, the prothorax and metathorax being much 
shorter. The mesonotum has its caudal margin produced into a long 
lobe, while in the other families the caudal margin of the mesonotum 
is very slightly curved. The appendages are very slightly soldered to 
each other and to the body, and the wings reach on to the fourth ab- 
dominal segment. The spiracles are small, circular, and very slightly 
produced. The pupae are from 6-8 mm. in length. The genera may 
be separated as follows : 

a. Body setae longer than the abdominal segments, heavily chitinized 
and forked at the end ; maxillae, measured on the meson, about half 
the length of the wings ; abdominal segments without deep punctures 
along the cephalic margin, but with a row of sharp triangular spines. 

BreniJiia Clemens. 

aa. Body with very short inconspicuous setae ; maxillae extending to the 

caudal margin of the wings ; abdominal segments 2-6 with a row of 

deep punctures along the cephalic margin, and with a row of 

sharp triangular spines just cephalad of the punctures. 

Choreutis Hiibner. 

The following species were examined : 
Brenthia pavonacella Clemens 
Choreutis inflatella Clemens, gnaphiella Kearfott 

SuPERFAMiLY AEGERIOIDEA 

The Aegerioidea, together with the Tortricoidea retain freedom of 
movement in abdominal segments 3-7 in the male and 3-6 in the 
female. The appendages are soldered to the body so that there is no 
ventral movement possible between the first two abdominal segments ; 
but there is undoubtedly dorsal movement, and at dehiscence these 
segments separate very distinctly from each other and the thorax, indi- 
cating that they have only recently lost their power of motion. In this 
superfamily is included the one family Aegeriidae. They form a very 
compact group in which it is hard to find satisfactory characters differ- 
entiating the genera. Moreover, pupae in good condition are difficult 
to obtain ; but it is hoped that the characters used here in separating 
the genera and in defining the superfamily will hold good for those 
groups to which they are applied. The sexes vary considerably and 
it has not been possible in all cases to obtain both male and female. 
This superfamily has most often been associated with the Tineoidea, 
but pupal characters indicate a much closer relationship to the Tortri- 
coidea. It is apparently somewhat nearer to the primitive famiHes 
Eriocraniidae and Nepticulidae than the Tortricoidea, owing to the 



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fact that a very large maxillary palpus is present in all genera, and that 
spines which reach well around to the ventral surface are found on 
abdominal segments 2-10, especially on the tenth segment. There 
are no setae yet developed on the anal rise, and there is not as much 
consolidation of the fixed caudal abdominal segments. The seventh 
segment in the female seems but recently to have lost power of motion. 
The abdominal segments are more nearly equal in length than in the 
Tortricoidea. 

Family Aegerddae 

The pupae of this family vary considerably in size, from the genus 
Aegeria, with species varying from 8-16 mm. in length, to the genera 
Memythrus and Bembecia, containing the largest species examined, 
varying from 20-25 mm. They are all provided with various forms 
of cutting plates for working their way to the surface, most of these 
being on the head, which is heavily chitinized at the cephalic end and 
usually has many ridges and projections, making it difficult to deter- 
mine the sutures. The clypeus often bears a sharp transverse ridge, 
sometimes toothed, which undoubtedly serves the same purpose. The 
body is elongate, cylindrical, with the abdominal segments approxi- 
mately equal in length, showing a generalized condition. The arrange- 
ment of parts in a pupa of this family is shown in Figures 36 and 37. 
It will be noted that the maxillary palpi are very large, and they remain 
uniformly so throughout the family. The appendages extend beyond 
the wings in most of the genera, but the caudal parts of these are not 
soldered to the body wall. The fronto-clypeal suture is always dis- 
tinct along the lateral margins of the front from the proximal ends of 
Ihe antennae almost to the invaginations for the anterior arms of the 
tentorium, but only shows transversely as a paler band of color in the 
strongly chitinized cuticle as indicated by the dotted line in Figure 36. 
Dehiscence invariably follows the course of this suture, and the front 
with the antennae are separated from the rest of the head parts. The 
epicranial suture is often obscured by the numerous elevations of the 
vertex and front, but it is always present. The antennae are always 
enlarged at the proximal end and again at the distal end, where they 
are somewhat club-shaped, thus differing again from the Tortricoidea. 
The mandibles are distinctly elevated in most genera. The wings are 
narrow and pointed, differing markedly from those of the Tortri- 
coidea. They are not soldered to the body wall at their distal end. 
The thorax always has a carinate median ridge, which may be distinct 
on all the segments, and is always distinct on the prothorax and the 
cephalic half of the mesothorax. The alar furrows are very deep, and 
one edge, usually the mesal one, is sharp and heavily chitinized. There 
are always two rows of spines on the dorsum of some of the abdom- 



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inal segments, which extend around to the ventral surface. These 
rows of spines are always present on segments 3-6, the number vary- 
ing on segments 2 and 7, while there is always one row on segments 
8-10. The number of rows of spines on segment 7 differs in the 
sexes, there being two rows in the male and only one in the female. 
The spines on the tenth segment are very broad, and this row extends 
nearly to the ventro-meson. Each of the spines has a seta inserted 
near its tip, which is not heavily chitinized and therefore easily broken. 
There are never any setae present on the anal rise. The genera of 
Aegeriidae may be separated as follows : 

a. Maxillae always more than half the length of the wings, generally 
nearly or quite equaling their length; eoxae of mesothoracic legs 
never adjacent on the meson below the maxillae, 
b. Clypeus with a prominent elevation near its caudal margin, bearing 
a heavily chitinized transverse ridge or series of projections which 
are probably to assist the pupa in cutting its way out of the 
burrow, 
c. Clypeus with the chitinized transverse ridge produced into a dis- 
tinct point on each side of the meson, 
d. Mesothorax with the median carinate ridge extending nearly 
its whole length, very distinct on the metathorax ; second ab- 
dominal segment with the two rows of spines distinct m both 
sexes ; maxHlae never reaching the caudal margin of the wings 
and ending opposite to the antennae ; pupae averaging 20-25 

mm. in length Sanninoidea Beutenmiiller. 

dd. Mesothorax with the median carinate ridge usually extending 
only along the cephalic half, never distinct on the meta- 
thorax; second abdominal segment never with two distinct 
rows of spines in either sex; maxillae reaching the caudal 
margin of the wings and ending opposite the mesothoracic 
legs; pupa usually 8-15 mm. in length. 

Sifncmthedon Hiibner. 
cc. Clypeus with a transverse row of separate projections. 

Parharmonia Beutenmiiller. 
bb. Clypeus not prominently elevated at its caudal margin and never ^ 
bearing ridges or projections which could be used m cutting, 
c. Tenth abdominal segment with eight spines m a row; caudal end 
of body just cephalad of the anal opening without setae. 

Podosesia Moschler. 

cc. Tenth abdominal segment with ten large spines in a row and two 

smaller ones, one on each side of the meson ; caudal end of body 

just cephalad of the anal opening with a row of four setae 

which are inserted under small projections. 

MemytJirus Newman, 
aa. Maxillae about two fifths the length of the wings; coxae of meso- 
thoracic legs and their tarsi adjacent on the meson caudad of the 
maxillae Bembecia Hiibner. 



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51 

The following species were examined : 
Sanninoidea exitiosa Say 
Synanthedon Hpuliformis Clerck, acerni Clemens, pictipes Grote and 

Robinson, pyri Harris, scitula Harris 
Parharmonia pirn Kellicott 
Podosesia syringae Harris 

Memythrus asilipennis Boisduval, dollii Neumoegen 
Bembecia marginata Harris 

SuPERFAMiLY TORTRICOIDEA 

This superf amily, like the Aegerioidea, is distinguished by the pres- 
ence of two rows of spines on the dorsum of most of the abdominal 
segments. The Tortricoidea form a more compact group than the 
Aegerioidea in regard to the arrangernent of appendages, which varies 
so little throughout the families that any member of the superfamily 
may be easily recognized by this arrangement, together with the pres- 
ence of spines on the abdominal segments. This characteristic arrange- 
ment is shown in Figures 38, 40, 41, and 44. There are often projec- 
tions from the head, much as in the Aegerioidea, but there are never as 
many head sutures present. The thorax shows the alar furrows in 
many instances but they are never as well developed as in the preced- 
ing superfamily, and never have sharp chitinized edges. The abdomen 
also shows a greater degree of specialization and its fixed caudal seg- 
ments are much more strongly consolidated, the sutures being very 
difficult to determine in many cases. The seventh segment has also 
become firmly fixed in the female. 

It was found impossible to group the pupae of this superfamily 
according to any of the schemes of classification now in use. The four 
groups into which the Tortricoidea discussed in the following pages 
have been divided are designated as Epiblemidae, Olethreutidae, Tor- 
tricidae, and Sparganothidae. These names, however, are without any 
significance whatever as far as previous classifications are concerned, 
and are merely used as a matter of convenience. Lack of material 
has prevented further study in this group at present, so it has been 
impossible to determine the correct family names. No attempt has 
been made to bring the nomenclature up to date. The generic names 
used by Meyrick and Walsingham have been followed as nearly as 
possible. 

The four groups or families of Tortricoidea must have had a 
common ancestor, but owing to the development of the maxillary pal- 
pus within the groups it would be impossible to consider one as 
derived from another. The line of development appears to have 



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been towards ( i ) a reduction of the spines on the dorsum of the ab- 
dominal segments, these disappearing first from the tenth segment and 
then from the segments cephalad of it; (2) the loss of setae on the 
anal rise; and (3) the development of a long cremaster. The families 
of Tortricoidea may be separated by the following table : 

a. Body without a distinct cremaster; setae always present on the anal 

rise Bpiblemidae. 

aa. Body with a well-developed cremaster. 

b. Ninth abdominal segment always with a distinct row of spines, 
especially in the males; tenth abdominal segment sometimes 
possessing spines ; cremaster broader than long ; setae always pres- 
ent on the anal rise, 
c. Cremaster never curved ventrad, the caudo-lateral angles not pro- 
duced into prominent hooks; the caudal margin usually showing 
three short lobe-like projections; second abdominal segment 
with the cephalic row of spines present and the caudal row well 
developed; setae of the anal rise always laterad of the anal 

opening Olethreutidae. 

cc. Cremaster curved ventrad, the caudo-lateral angles produced 
into prominent hooks ; second abdominal segment lacking the 
cephalic row of spines and the caudal row poorly developed ; 
setae of the anal rise always on the caudal part of the eleva- 
tion TORTRICIDAE. 

bb. Ninth abdominal segment lacking a distinct row of spines, although 
a few spines are sometimes present in the males; setae never 
present on the anal rise; cremaster nearly always longer than 
broad ; tenth abdominal segment never possessing spines. 

Sparganothidae. 

Family EpiBLEMroAE 

The pupae belonging to this family (Figs. 38, 39) have no cre- 
master and there are always setae present on the anal rise. They are 
usually less than 10 mm. in length and slender, tapering gradually 
from the thoracic region to the somewhat blunt end of the body. The 
genus Carpocapsa is sometimes an exception as the body is often very 
stout, and the genus Eucosma has a cyHndrical body strongly resem- 
bling the pupae of the Aegeriidae. The maxillary palpi usually extend 
to the proximo-lateral angles of the maxillae; only Epinotia and 
Enarmonia of the genera studied had shorter palpi. The maxillae are 
about two fifths the length of the wings, and the labial palpi are usually 
half the length of the maxillae. The rows of spines on the dorsum 
vary somewhat in the different genera. All have two rows present on 
abdominal segments 2-7 although the cephaUc row of segment two is 
weak in Eucosma, Hemimene, and some species of Ancylis. Occa- 



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sionally the caudal row of segment seven is weak in the females of 
some species. As a general rule the eighth segment has but one row 
of spines, the cephalic one, but two rows have been found in the species 
of Epinotia and Eucosma, usually in the males. There is, in most 
genera, considerable difference between the sexes. The antennae do 
not vary as greatly in this family as in some others, but there is a 
great variation in the rows of spines, these being usually smaller and 
less numerous on the caudal row of segment seven in the male and 
on segment eight in the female. The genus Mellisopus does not show 
characters of sufficient importance to allow of its retention as a sepa- 
rate genus, and it is therefore included with Carpocapsa. The only 
points of difference between that genus and Carpocapsa pomonella, 
its nearest ally, are that the spiracles are oval, somewhat rectangular 
and slightly produced, while in Mellisopus latiferreanus they are large 
and circular but not strongly produced. There is however consider- 
able variation. Mellisopus shows a slight carinate ridge on the meta- 
thorax, but this, again, is extremely variable. 

The phylogeny of the group is extremely doubtful. If the spines 
on the dorsum of the abdominal segments in Eucosma were homol- 
ogous with those found in the generalized pupae of Nepticulidae and 
others, it would certainly be the most generalized form and the others 
would probably follow the sequence of the table to genera. The 
spines, however, are much broader than any observed in the general- 
ized types. Eucosma also shows a remarkable resemblance to the 
Aegeriidae, so it is probable that it is the most generalized of the 
Tortricoidea examined. The genera of Epiblemidae may be separated 
a^ follows : 

a. With two long distinct setae present on each side of the anal rise, 
b. Caudal end of body with one row of long, heavily chitinized, flat- 
tened setae inserted along the line of the row of spines on the 
tenth abdominal segment, 
c. Dorsal surface of abdominal segments between the cephalic and 
caudal rows of spines covered, at least in part, with short trian- 
gular spines; cephalic row of spines composed of alternating 

large and small spines Eucosma Httbner. 

cc. Dorsal surface of abdominal segments between the caudal and 

cephalic rows of spines always smooth ; cephalic row of spines 

of approximately the same size. 

d. Portion of first coxae exposed on the meson below the maxillae 

more than half the length of the second coxae; body often 

stout, with the length scarcely three times the greatest width, 

but extremely variable ; length averaging 10 mm. 

Carpocapsa Treitschke. 



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dd. Portion of first coxae exposed on the meson below the maxillae 
never more than half the length of the second coxae ; body 
always slender, with the length about four times the greatest 

width ; average length 8 mm Tmetocera Lederer. 

bb. Caudal end of body with two rows of setae showinsr. one row of 
four setae inserted along the line of the row of spines on the tenth 
abdominal segment and another row at the caudal margin of 
the body, 
c. Caudal row consisting of four setae; maxillary palpi always 
touching the proximo-lateral angles of the maxillae, 
d. Caudal row of setae of two kinds, the mesal ones much slen- 
derer than the lateral ones ; maxillae less than one third the 
length of the wings; labial palpi about two thirds the length 
of the maxillae ; maxillary palpus touched by the prothoracic 

leg Hemimene Hiibner. 

dd. Caudal row of setae similar; maxillae never less than one 
third the length of the wings; labial palpi about half the 
length of the maxillae ; maxillary palpus touched by both 

prothoracic and mesothoracic legs Ancylis Hiibner. 

cc. Caudal row consisting of two setae; maxillaiy palpi seldom 
reaching the proximo-lateral angles of the maxillae. 

Epinotia Httbner. 

aa. Never with two long, distinct setae on each side of the anal rise. 

b. Lateral spines of the tenth row noticeably larger than the others; 

setae at the caudal end of the body very short and slender, not 

heavily chitinized ; setae of the anal rise very small and difficult 

to locate, usually two present on each side Thiodia Httbner. 

bb. Lateral spines of the tenth row not noticeably larger than the 
others ; setae at caudal end of body long and heavily chitinized ; 
one seta on a distinct papilla on each side of the anal rise. 

Enarmonia Httbner. 
The following species were examined : 
Eucosma strenuana Walker, scudderiana Clemens 
Carpocapsa pomonella Linnaeus, saltitans Westwood, latiferreanus 

Walsingham 
Tmetocera ocellana Schiflfermueller 
Hemimene incanana Clemens 

Ancylis comptana Frolich, platanana Clemens, diminutana Kearfott 
Epinotia saliciana Clemens, piceafoliana Kearfott 
Thiodia signatana Clemens 
Enarmonia fana Kearfott 

Family Olethreutidae 

The Olethreutidae (Fig. 40) include those species which possess a 
well-developed cremaster, usually broader than long and somewhat 



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flattened, bearing eight strongly chitinized, flattened, hooked setae ; and 
usually having similar but smaller setae on the anal rise. Exceptions 
to this latter character are found in the genus Polychrosis, and in 
Bxartema ferriferanum, which does not agree with the remainder of 
the genus in this respect. The group is further characterized by the 
presence of a well-developed row of spines on the ninth abdominal 
segment in all the males examined and in nearly all of the females, the 
exceptions being in the genus Exartema, where the spines were smaller 
and fewer in number. In most genera this row of spines has several 
additional spines on each side, usually near the meson. The only 
other species of the superfamily which resemble the members of this 
group are the species of Archips in group (b), but these have no setae 
on the anal rise, and very seldom have spines present on the ninth 
abdominal segment. Exartema ferriferanum is the only species among 
those examined which might be confused, as the row of spines on 
the ninth segment of the female is not well developed, while the males 
of Archips cerasivorana sometimes have a few spines present. This 
particular species of Exartema, however, has a prominent cephalic 
projection, ending in a point, directed ventrad, while the species of 
Archips are blunt at the cephalic end, and the bodies are usually larger 
and prominently enlarged in the region of the thorax. The antennae 
show marked sexual differences, being much longer in the males. The 
rows of spines on the dorsum of the abdominal segments also vary in 
the sexes, the caudal row of segment eight being poorly developed or 
lacking in many females, though well developed in the males. The 
row on the ninth segment is much better developed in the males. The 
genus Polychrosis shows a peculiar development of the spiracles. 
Instead of the small, produced tubular spiracles common to the Tortri- 
coidea it appears to have them very much enlarged. This prominent 
enlargement around the sprtracle has a deeply concave surface, and the 
very small tubular spiracle in the center is about one sixth of its width. 
A similar condition, but not so well developed, is found in Exartema 
sciotoanum. The maxillary palpi are well developed and reach the 
proximo-lateral angles of the maxillae in Olethreutes (b) and Poly- 
chrosis, but in Episimus, Olethreutes (a), and Exartema they are not 
well developed. The genera of Olethreutidae may be separated as 
follows : 

a. Tenth abdominal segment with spines, usnaUy three or four rows 

closely approximated, seldom with a single row. 

b. Long chitinized setae present on the anal rise, usuaUy slightly 

shorter and narrower than those of the eremaster. 

c. With two setae on each side of the anal rise, very similar to those 

on the eremaster Episim^is Walsingham. 



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cc. With one seta on each side of the anal rise, smaller than those 

of the cremaster Olethreutes (a) Hubner. 

bb. Setae never present on the anal rise PolycJirosis Ragonot. 

aa. Tenth abdominal segment without spines, 
b. Well-developed setae present on each side of the anal rise. 

e. Maxillary palpi well developed, reaching the proximo-lateral 

angles of the maxillae Olethreutes (b) Hubner. 

cc. Maxillary palpi short, never reaching the proximo-lateral angles 

of the maxillae * Exartema (a) Clemens. 

bb. Without setae on the anal rise Exartema (b) Clemens. 

The following species were examined : 
Episimus argutanus Clemens 

Olethreutes (a) niveiguttana Grote, (b) malachitana Zeller 
Polychrosis sling erlandana Kearfott, viteana Clemens, botrana Schif- 

fermueller 
Exartema (a) sciotoanum Kearfott, concinnanum Clemens, nigra- 

num Kearfott, inornatum Clemens, permundanum Clemens 
Exartema (b) ferriferanum Walker 

Family Tortricidae 

This group is distinguished by its peculiar type of cremaster and 
the presence of setae on the anal rise. The maxillary palpi are not 
present in Peronea but are found in Argyrotoxa, where they are 
shorter in the male than in the female. The maxillae are usually about 
two fifths the length of the wings, the labial palpi nearly half the 
length of the maxillae. There are no spines present on the tenth ab- 
dominal segment, and they are not well developed on the second and 
third segments. There is always a well-developed cephalic row on the 
dorsum of the tenth segment, but the caudal one does not extend as far 
laterad in the male and is usually lacking in the female. In Argyrotoxa 
the cephalic row of spines on the eighth and ninth segments is on a 
prominent ridge which can be plainly seen on the lateral margin in 
dorsal view. There are always two setae present on each side of the 
anal rise and these are always on the caudal part of the elevation. 
Figures 41 and 42 show the arrangement of parts in this family and 
Figure 43 the dehiscence of part of the head, showing the eye-piece. 
The genera of Tortricidae may be separated by the following table : 

a. Maxillary palpi present in both sexes ; spines of the cephalic row on 
abdominal segments 7-9 on distinct ridges which show plainly on the 
lateral margins of the body, the spines extending laterad beyond the 
spiracles in some segments; setae on the ventral side of cremaster 
and between cremastral hooks not heavily chitinized and resembling 
the ordinary body setae Argyrotoxa Stephens. 



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aa. Maxillary palpi not present in either sex ; spines of the cephalic row 
on abdominal segments 7-9 not on distinct ridges, the spines never 
extending laterad beyond the spiracles ; setae on the ventral side of 
cremaster heavily cWtinized, and usually not extending far beyond 
the caudal margin of the body Peronea Curtis. 

The following species were examined: 
Argyrotoxa albicomana Clemens, bergmanniana Linnaeus 
Peronea sp., tninuta Robinson, logiana Schiflfermueller, var. viburnana 
Clemens 

Family Sparganothtoae 

This family (Fig. 44) includes the species in which the cremaster 
is well developed and much longer than broad, except in Archips (b) 
and Phaecasiophora. The cremaster in nearly all species bears eight 
strong hooked setae which are usually not much flattened except in 
the genera mentioned above. There are never any setae present on 
the anal rise, and most 6f the species have no spines present on the 
ninth abdominal segment and none of them a well-developed row. The 
caudal row of the eighth segment is often lacking in the female and 
is poorly developed in the male. The females of Platynota flavedana 
have no cephalic row on the second abdominal segment. The members 
of this group include the largest of the Tortricoidea examined, most 
of them considerably over 10 mm. in length; the thoracic region 
usually appears considerably enlarged, and the abdomen is long and 
tapering. The vertex is shorter than in the other groups. The max- 
illary palpi do not reach the proximo-lateral angles of the maxillae in 
the males, but sometimes do so in the females. In Platynota flavedana 
the palpi appear to extend only along the cephalic margin of the pro- 
thoracic leg. The setae of the body are usually very long and promi- 
nent in this group. Sexual differences are noticed in the length of 
the maxillary palpi and antennae and in the development of the rows 
of spines on the dorsum of the abdominal segments. There are no 
available characters by which all the species of the genus Archips can 
be associated in a single group and it undoubtedly represents two 
genera, because there are two distinct types' of cremaster present. It 
is also difficult to find good structural characters to separate the genera 
Harmologa and Archips (a). The color markings are very distinct in 
Harmologa, and the body is also very noticeably enlarged in the region 
of the first three abdominal segments, so that in ventral view the lateral 
margins of the wings appear curved, instead of approximately parallel 
as in Archips (a). The genera Epagoge and Platynota are also closely 
related and are grouped together by some writers. The genera of 
Sparganothidae may be separated as follows : 



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a. Transverse conjunctiva showing prominent dark brown spines scat- 
tered over a lighter brown surface, 
b. Cremaster much longer than broad, not flattened. 

c. With four setae inserted at the caudal end of the cremaster. 
d. Dorsum of second abdominal segment showing a slightly crenu- 
late, chitinized cephalic margin, the cephalic row of spines on 
this segment not well developed in the males and wanting in 
females; head never with a cephalic projection; abdomen 
never with prominent cavities on the dorsum of the second 
and third segments ; dorsum of abdomen always with darker 

transverse bands and spots of color Harmologa Meyrick. 

dd. Dorsum of second abdominal segment without a crenulate 
cephalic margin; cephalic row of spines well developed on 
this segment in both sexes ; head often with a cephalic pro- 
jection, or if not, then prominent cavities are present on the 
dorsum of the second and third abdominal segments; body 

of uniform color Archips (a) Hiibner. 

cc. With two setae inserted at the caudal end of the cremaster. 

Cenopis Zeller. 

bb. Cremaster broader than long, flattened Pliaecasiophora Grote. 

aa. Transverse conjunctiva never showing prominent dark brown spines, 
surface of uniform color, 
b. Cremaster longer than broad, not flattened; labial palpi always 
considerably more than half the length of the maxillae, 
c. Cephalic row of spines on second abdominal segment lacking in 
the female ; cremastral setae noticeably flattened. 

Platynota Clemens, 
cc. Cephalic row of spines on the second abdominal segment present 
in the female ; cremastral setae not noticeably flattened. 

Epagoge Hubner. 

bb. Cremaster broader than long, distinctly flattened ; labial palpi not 

more than half the length of the maxillae. .Archips (b) Hubner. 

The following species were examined : 

Harmologa fumiferana Clemens 

Archips (a) argyrospila Walker, magnoliana FtrnsAdjparallela Robin- 
son, obsoletana Walker, rosaceana Harris 

Archips (b) cerasivorana Fitch, fervidana Clemens 

Cenopis chambersana Kearf ott 

Phaecasiophora confixana Walker 

Platynota flavedana Clemens 

Epagoge sulfureana Clemens 

SuPERPAMiLY GRACILAMOIDBA 

This superfamily name is given to a number of families apparently 
of common origin, which have proceeded along similar lines of devel- 



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59 

opment. The species are all leaf -miners and are very small, the pupae 
of the largest species examined being 7 mm. in length. Very few of 
the generalized families have been available for study, so that it is 
exceedingly difficult to trace the relationships existing between the 
more specialized families without first having carefully studied a num- 
ber of more generalized forms. There is included in this group the 
Nepticulidae, in many respects the most generalized pupae studied, 
next to the Eriocraniidae, and certainly resembling the latter more 
than any of the other generalized forms examined. It is just at this 
point in our investigation that more material is needed to clear up the 
relationships of the groups which have apparently branched off here 
and have had a common ancestor with the Nepticulidae. From all the 
evidence at hand it seems probable that development has proceeded 
along two well-defined lines, the first, represented by the superfamily 
Gracilarioidea, having early lost the maxillary palpi while still retain- 
ing the covering of spines on the dorsum of the abdominal segments, 
and having developed the triangular type of prothorax; the second 
having retained the maxillary palpi for a much longer time, but having 
lost the covering of spines, while developing the same type of pro- 
thorax. 

Of the second type no material has been examined which would 
show any intermediate stages between the families Nepticulidae and 
Epermeniidae. The latter family has apparently continued the line 
of development begun in the Gracilarioidea as it still retains the 
seventh abdominal segment free in the male though it is fixed in the 
female. The presence of the maxillary palpi precludes its derivation 
from the Gracilarioidea and would lead us back to some point below 
the Heliozelidae because this family also has lost them. As we have 
only the Nepticulidae for comparison, it has been assumed that this 
branch has arisen coordinately with them. 

In the superfamily Gracilarioidea, with the exception of the family 
Lyonetiidae, all the pupae have free appendages, the cuticle is very 
slightly chitinized, and the dorsum of the abdomen is covered, in part 
at least, with fine spines. There is a tendency in some genera, as 
LithocoUetis and Ornix, towards the development of a single row of 
spines, so that there is often one or more rows of larger spines at 
either the cephalic or caudal margin, or at both margins, of the seg- 
ment. This seems to indicate the way in which the rows of spines were 
developed in the Tineoidea and Tortricoidea. The characters which 
are common to all the members of the superfamily are the long vertex, 
which is always longer than the prothorax at the median line, scarcely 
ever less than twice its length and often much longer, and the long 



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metathorax, with the loss of a well-developed maxillary palpus in 
families above the Nepticulidae. Chapman described the genus Graci- 
laria as possessing maxillary palpi, and in two species, sassafrasella 
and negundella, a structure has been found (Figs. 45, 47) which may 
be the maxillary palpus ; but there never is a distinct, oblong piece lying 
caudad of the eye-piece as is usually the case when the maxillary palpi 
are present, and of all the species of the superfamily examined these 
two were the only ones in which there was any doubt as to its absence. 
The head is in most families either produced into a prominent projec- 
tion or there is a heavily chitinized cutting plate near the cephalic mar- 
gin on the ventral surface. The prothorax has a tendency to become 
shorter on the median line and longer on its lateral margins, so that 
each half is triangular. In such cases the length along the lateral 
margin is about four times the length on the median line. In the more 
generalized forms the prothorax is more like the rectangular type 
found in the Tineoidea, but it is depressed or sunken, giving it a neck- 
like appearance. The metathorax still retains its primitive condition, 
and is usually more than half the length of the mesothorax. In nearly 
all of the families the wings are long in proportion to the body, and in 
the majority they are about two thirds its length. The bodies of most 
of the families included here retain the generalized type found in the 
Eriocraniidae with a slight depression near each lateral margin in the 
region of the spiracles. The spiracles are usually small, circular, and 
slightly produced, appearing tubular. The Lyonetiidae seem to be an 
exception to almost every rule. They have no free segments, the 
appendages are all soldered to the body, and there are no spines visible 
on the abdomen. They seem to be more nearly related to the Buc- 
culatrigidae than to any other family, although there are strong rea- 
sons for considering them related to the Phyllocnistidae. The follow- 
ing table will serve to separate the families of Gracilarioidea : 

a. Maxillary palpi well developed and extending along the caudal margin 
of the eye ; spiracles visible on the first abdominal segment. 

Nepticulidae. 

aa. Maxillary palpi never well developed, and if present never extending 

as an oblong piece along the caudal margin of the eye. 

b. Antennae never extending more than half the length of the wings ; 

labrum very long and lobe-like, extending down over the labial 

palpi for about one fourth of their length ; spines on the dorsum 

of the abdomen very fine and not easily distinguished. 

Heliozeljdae. 

bb. Antennae always extending at least three fourths the length of the 

wings, and usually equaling them in length or extending beyond 

their caudal margin ; labrum never long and lobe-like and never 



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extending down over the labial palpi for one fourth of their 
length, 
c. Appendages free, never firmly soldered to the body wall ; abdomen 
always with some of the segments movable ; dorsum of the abdo- 
men always with spines, 
d. Abdominal segments 3-7 movable in the male, 3-6 in the 
female ; antennae and metathoracic legs not approximately 
equal in length and both seldom extending beyond the caudal 
margin of the wings, 
e. Labial palpi present ; caudal end of body ending in two stout 
spines directed dorsad; abdominal segments 3-6 with the 
two setae nearest the meson on the cephalic half of the seg- 
ment so closely approximated that their bases touch. 

TiSCHERIIDAE. 

ee. Labial palpi never visible ; caudal end of body never with 
curved hooks, but the tenth abdominal segment with a 
prominent lateral projection on each side ending in a stout 
straight spine; abdominal segments 3-6 with the setae 
nearest the meson at the cephalic end of the segment never 

closely approximated BuccuLATRioroAE. 

dd. Abdominal segments 4-7 movable in the male, 4-6 in the 
female; antennae and metathoracic legs approximately of 
equal length and both always extending beyond caudal mar- 
gin of the wings, 
e. Abdominal segments 3-7 never with two deep punctures or 
pits with heavily chitinized edges on the meson at the 
cephalic margin with one or more heavily chitinized spines 
adjacent; the l^gth of abdominal segments 8-10 always 

greater than that of segment 7 Gracilarhdae. 

ee. Abdominal segments 3-7 with two deep punctures on the 
meson at the cephalic margin with one to three heavily 
chitinized spines adjacent ; abdominal segments 8-10 never 
showing distinct segmentation, their total length less than 

that of the seventh segment Phyllocnistidae. 

cc. Appendages always firmly soldered to the body ; dorsum of abdo- 
men without visible spines; abdomen without any movable seg- 
ments Lyonetiidae. 

Family Nepticuudae 

These tiny species of leaf-miners average 2 mm. in length in the 
females and 1.5 mm. in the males. The body is flattened, with a trans- 
parent, slightly chitinized cuticle, and is white in color until the adult 
scales are formed. Although their size makes it difficult to determine 
the number of free segments, it is believed that there is some degree 
of motion between all of the abdominal segments except the fixed 



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caudal ones. There is some degree of movement between the seventh 
and eighth abdominal segments in both sexes, but it is apparently 
greater in the male. The arrangement of parts may be seen in 
Figures 48 and 49. The head does not show all of the sutures found 
in the Eriocraniidae, but the epicranial and fronto-clypeal sutures are 
always present. The appendages are all free and segmented as in the 
Eriocraniidae, and the thoracic appendages are widely separated to 
show all the coxae. There is a strong resemblance between this family 
and the more generalized members of the Eucleoidea, but the presence 
of the large maxillary palpi prevents their being included in that super- 
family. The spiracles are visible on the first abdominal segment, and 
the length of the thoracic segments indicates a very generalized condi- 
tion. The genital opening of the male is located as shown in Figure 
48. In the females there is an area covered with setae on the venter 
of the eighth segment, as in the Eriocraniidae, but no openings could 
be accurately determined. 

The following species were examined : 
NepHcula nyssaefoliella Chambers, platanella Clemens. 

Family Heuozelidae 

This family includes some very small pupae which measure only 
2-3 mm. in length. They have all the appendages free and widely 
separated. The cuticle is transparent and the body white in color, with 
the conjunctiva so little differentiated that it was impossible to deter- 
mine the number of free segments with accuracy. Segments 2-7 in 
the male and 2-8 in the female have some power of motion, but 
whether this is movement of the whole segment in the case of the 
second and third, or merely dorsal movement, was not determined. 
The family (Fig. 50) is characterized by its short antennae, and its 
long labrum which projects down over the labial palpi. They also 
have shorter appendages than any of the other families with transpar- 
ent cuticle and white bodies, because in all others the metathoracic 
legs and antennae extend considerably beyond the caudal margin of 
the wings and are often longer than the body. The epicranial suture 
is near the cephalic margin of the head. While this family may have 
retained more free segments than the Gracilariidae it is undoubtedly 
more specialized than some of the genera in that family. The pro- 
thorax is much longer at its lateral margins than on the meson ; there 
is no trace of maxillary palpi, and the labial palpi are not so well 
developed as in the generalized Gracilariidae. 

The genera included in this family have long been associated with 
the Elachistidae, but the pupae show no resemblance whatever to this 



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68 

family. The name Heliodinidae has been applied by some authors to 
the genera included in this family, but Meyrick, in Lepidoptorum 
Catalogus, Part 13, uses this name to include the genera Brenthia, 
Choreutis, etc. The name Heliozelidae is used by Spuler (Die Schmet- 
terlinge Europas, 1910) and this name has been adopted there. The 
genera may be separated as follows : 

a. Abdomen with one or two prominent lateral setae on each side of the 
tenth abdominal segment ; mesonotum not produced into a prominent 

lobe extending down on the metathorax Aniispila Hubner. 

aa. Abdomen never with prominent lateral setae on each side of the tenth 
abdominal segment; mesonotum produced into a prominent lobe 
extending down on the metathorax Coptodisca Walsingham. 

The following species were examined : 
Antispila ampelopsisella Chambers, cortdfoliella Clemens 
Coptodisca juglandiella Chambers, splendiforella Clemens 

Family Tischerhdae 

These pupae are from 3.5-6 mm. in length and have abdominal 
segments 3-7 free in the male and 3-6 in the female. They are always 
considerably chitinized, so that the pupae vary in color from yellow 
to brown. The spines on the dorsum of the abdominal segments are 
very distinct and in some species they are of two sizes. All of the 
species examined except Tischeria heliopsisella (Fig. 54) had certain 
of the body setae very long, heavily chitinized, and forked at the end. 
These setae vary in length, but the shortest are nearly as long as the 
abdominal segments and are very conspicuous. The dorso-mesal setae 
nearest the cephalic margin were closely approximated on segments 
3-6 or 7 of the abdomen so that their bases were in contact. The 
caudal end of the abdomen is bifurcate, and ends in two heavily chitin- 
ized hooks which are directed dorsad. The arrangement of parts may 
be seen in Figures 51-54. These pupae have become more specialized 
in certain respects than many of the Gracilariidae, although they re- 
tain one more free segment. This is noticeable in the development of 
the prothorax, in the distinct rows of larger spines on many of the 
segments, and in the strong caudal hooks. The f ronto-clypeal suture 
shows as a clear area, indicated by the dotted line in Figure 51. This 
family includes two genera, Coptotriche and Tischeria, with no well- 
defined characters for separating them. The two species of Tischeria, 
aenea from blackberry and malifoliella from apple, at one time consid- 
ered identical, show distinct differences in the pupae and both species 
resemble Coptotriche, while heliopsisella is very different from all the 



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64 

rest. Dyar's list names but one species of Coptotriche, but three dis- 
tinct types of pupae have been obtained from mines in oak leaves. 
Unfortunately no adults have yet emerged from these, so the species 
can not be determined. The genera may be separated as follows : 

a. Caudal margin of the dorsum of the second abdominal segment heavily 
chitinized and toothed, the teeth being larger than the adjoining 
spines Coptotriche Walsingham. 

aa. Caudal margin of the dorsum of the second abdominal segment not 
heavily chitinized or toothed Tischeria Zeller. 

The following species were examined : 
Coptotriche zelleriella Clemens 

Tischeria aenea Frey and Boll, malifoliella Clemens, heliopsisella 
Chambers 

Family BuccuLATRiGroAE 

This family, Bucculatrigidae, including the single genus Buccu- 
latrix, has been placed in various positions by different authors. It 
is quite evident that it is more specialized than most other families of 
the Gracilarioidea in the loss of the labial palpi and that it has pro- 
ceeded along a different line of development. Nevertheless, no one can 
fail to see the relationship between the pupae of the Bucculatrigidae 
and the other members of this superf amily, particularly to some of the 
species of Cameraria where there is a lateral projection from each side 
of the tenth segment and a distinct row of larger spines on the dorsum 
of the abdominal segments. The lack of labial palpi, together with the 
spines on the abdominal segments, is sufficient to distinguish the 
family from all the others included in the superfamily. The arrange- 
ment of parts may be seen in Figures 55 and 56. The pupae examined 
had an average length of 3 mm. 

The following species were examined : 
Bucculatrix sp., pomifoliella Clemens, trifasciella Clemens. 

Family Lyonetodae 

This family is a very difficult one to place satisfactorily by pupal 
characters alone, as it has completely lost the power of motion in the 
abdominal segments and all the appendages are soldered down. This 
is another of the families which has been a source of anxiety to many 
lepidopterists. The shape of the prothorax, the length of the vertex, 
together with that of the wings and appendages as compared with the 
body, the small tubular spiracles, and the absence of maxillary palpi 
seem without any doubt to indicate its relationship to the members 
of the superfamily Gracilarioidea and consequently it is included here. 



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65 

From a careful study of the pupal characters available it seems to be 
more nearly related to the Bucculatrigidae than to any other family. 
A comparison of Figure 57 or 59 with 67 will show that the develop- 
ment in the Lyonetiidae has not been towards the shortening of the 
segments and the consolidation of abdominal segments 8-10 as in 
the Phyllocnistidae. Moreover, it still retains the generalized type 
of body found in the Nepticulidae, while the Phyllocnistidae have 
developed the cylindrical type. The shape of the maxillae and the 
position of the femur of the prothoracic leg are as in the Bucculatrig- 
idae, and like them the Lyonetiidae have no labial palpi visible. The 
Lyonetiidae do not spend their pupal life within the mine, nor in a 
cocoon, but are exposed and fixed by the caudal end to some cross 
threads on the under surface of the leaf (Clemens, Tineina of N. 
America, 1872, pp. 189-191 ). The soldering down of the appendages 
and the loss of motion of the abdominal segments seems to be a modi- 
fication to suit the new conditions of life, and is analogous to the con- 
dition found in certain families of Papilionoidea and the species of 
the genus Elachista, in which all power of motion is lost. Bedellia 
has developed certain ridges and projections, similar to those found 
in the Papilionoidea, which seem to be correlated with this manner 
of pupal life. Only two genera of Lyonetiidae were studied. These 
were from 4-6 mm. in length and may be separated as follows : 

a. Head blunt, without a promment projection; antennae and meta- 
thoracic legs equal in length ; caudal end of body with a few straight 
spines on the dorsal surface of the tenth abdominal segment ; body 
without prominent ridges Proleucoptera Busck. 

aa. Head with a long projection ; antennae much longer than the meta- 
thoracic legs; caudal end of body with hooked setae; body with 
prominent ridges Bedellia Stainton. 

The following species were examined : 
Proleiuoptera smilaciella Busck 
Bedellia somnulentella Zeller 

Family Oracilaioidae 

This large family includes those pupae with free appendages and 
with abdominal segments 4-7 free in the male and 4-6 in the female. 
The antennae and metathoracic legs are of approximately the same 
length and both are longer than the wings. The most nearly related 
family, the Phyllocnistidae, differs in having on the dorsum of each 
abdominal segment two prominent pits or punctures with heavily 
chitinized edges associated with some large curved spines, in having 



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66 

a much more cylindrical body with large deep furrows between the 
segments, and in having the fixed caudal segments very short. 

The genus Gracilaria is undoubtedly the most generalized, if we 
consider the peculiar structures (Fig. 47) found in some species to 
be maxillary palpi. The tendency in the Gracilariidae is toward the 
loss of the maxillary palpi, and the development of the triangular 
type of prothorax, which usually is elevated on the median line. There 
is also a shortening of the maxillae and labial palpi and of all the ap- 
pendages in relation to the rest of the body, and a stronger chitiniza- 
tion of the surface of the body tending to a soldering down of the 
appendages. There is also taking place the development of two sizes 
of spines on the dorsum of the abdominal segments and the formation 
of single rows of larger spines. Finally there is the development of 
the cremaster. There are two distinct divisions of the Gracilariidae 
to which subfamily names have been given. These may be separated 
as follows: 

a. Prothorax depressed and neck-like, somewhat quadrangular in out- 
line, the length at the lateral margin never more than twice the 

mesal length Grachlariinae. 

aa. Prothorax usually with an elevated ridge on the meson, triangular 
in outline, the length at the lateral margin about four times the 
mesal length Lithocolletinae. 

Subfamily Gracilariinae 

The Gracilariinae (Figs. 45 and 46) include all the genera in 
which the generalized quadrangular type of prothorax has been re- 
tained. In all the genera the caudal end of the body is blunt and the 
tenth segment bears a row of 6 or 8 spines, larger than those on the 
other body segments. The labial palpi are always long and never 
covered by the maxillae at their proximal end. The following table 
will serve to separate the genera of Gracilariinae : 

a. Dorsum of abdomen sparsely covered with very coarse spines, some- 
times with additional fine spines, 
b. Head with a cutting plate on the ventral surface near the cephalic 
margin, which is usually serrate ; maxillae as long as the mesotho- 

racic legs GracUaria Haworth. 

bb. Head with a prominent projection at the cephalic end, not a 
distinct plate; maxillae never as long as the mesothoracic legs. 

Ornix Treitschke. 

aa. Dorsum of abdomen thickly covered with very fine spines which are 

almost invisible Parectopa Clemens. 



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67 

Subfamily Lithocolletinae 

In the Lithocolletinae all the genera but Acrocercops have a 
strongly elevated median ridge on the prothorax, and in all but Acro- 
cercops and Marmara the proximal part of the labial palpi is covered by 
the maxillae so that the lateral margin can not be traced cephalad to 
the labrum. The genus LithocoUetis, which seems very distinct from 
other genera in the subfamily, includes two distinct types of larvae. 
On this basis the genus was divided into two groups designated as 
the "flat-larval group" and the "cylindrical-larval group." Dr. Chap- 
man in 1902 (Entomologist, Vol. 35, p. 141) proposed the name 
Cameraria for the flat-larval group, and this name is used here as our 
investigation shows that the cremaster is a decided genus character, 
and, furthermore, that members of the same genus have the same type 
of cremaster. It is therefore deemed impossible, from a study of the 
pupal characters, that one genus could include both forms with and 
without a cremaster. The pupae of the cylindrical-larval group stud- 
ied, moreover, showed two distinct types of cremaster, L. lucidicos- 
tella (Fig. 66a) having a rather broad cremaster with curved setae, 
while in L. Hliacella and L. argentinotella the cremaster is long and 
slender (Figs. 66b and 64) and the setae are T-shaped, the former 
having one such seta and the latter two. From the standpoint of 
pupal characters these would properly form three genera. It is in- 
teresting to note that Meyrick (Genera Insectorum, Part 128) places 
these in different sections of the genus, and that Miss A. F. Braun 
in her work on the "Development of the Color Pattern in Lithocol- 
letis" (Journ. Acad. Nat. Sci. Phila., Vol. 16, Series 2, 1914) also 
includes them as members of different groups in her phylogenetic tree. 
The genera of Lithocolletinae may be separated as follows: 

a. Dorsum of abdominal segments with spines of the same size; caudal 
margins of abdominal segments never distinctly elevated, 
b. Dorsum of each abdominal segment covered with spines for its 
entire length, 
e. Maxillae at least seven eighths the length of the wings; labial 
palpi almost half the length of the maxillae, their proximal 
end not covered by the maxillae; spines on dorsum of abdo- 
men very small and inconspicuous except a row of six spines 

on the tenth segment Acrocercops Wallengren. 

cc. Maxillae not more than one third the length of the wings ; 
labial palpi about one third the length of the maxillae, their 
proximal end covered by the maxillae ; spines on the dorsum 
of the abdomen small but distinct, with a few larger ones on 
the tenth segment Leucanthiza Clemens. 



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68 

bb. Dorsum of each abdominal segment covered with spines for about 
one fourth its length, 
c. Head without a prominent pointed projection; maxillae more 
than half the length of the wings and longer than the pro- 
thoracic legs; proximal part of the labial palpi not covered 

by the maxillae* Marmara Clemens. 

cc. Head with a pVominent projection ; maxillae never half the 
length of the wings nor as long as the prothoracic legs ; labial 
palpi covered by the maxillae at the proximal end. 

Cremastobombycia Braun. 
aa. Dorsum of abdominal segments covered with spines of two sizes, the 
caudal margins of the segments usually distinctly elevated ; labial 
palpi always covered by the maxillae at the proximal end. 
b. Caudal end of body never with a distinct cremaster. 

Cameraria Chapman, 
bb. Caudal end of body always with a distinct cremaster. 

Lithocolletis Hiibner. 
Species of Gracilariidae examined : 
Subfamily Gracilariinae 

Gracilaria negundella Chambers, sassafrasella Chambers, violacclla 

Clemens 
Ornix prunivorella Chambers, crataegifoliella Clemens, conspic- 

uella Dietz 
Parectopa salicifoliella Chambers, lespedesaefoliella Clemens 
Subfamily Lithocolletinae 
Acrocercops venustella Clemens 

Leucanthiza amphicarpeaefoliella Clemens, ostensackenella Fitch 
Marmara salictella Clemens 
Cremastobombycia solidaginis Frey and Boll 
Cameraria hamadryadella Clemens, ostryella Chambers, tubiferella 

Clemens 
Lithocolletis lucidicostella Clem., argentinotella Clemens, tiliacella 
Chambers 

Family Phyllocnisttoae 

This family is very nearly related to the Gracilariidae, and the 
principal characters used to distinguish it are given under that family. 
The arrangement of parts may be seen in Figure 67. It will be noted 
that Phyllocnistis has long heavily-chitinized setae much as in the 
Tischeriidae except that they are not forked at the tip. There is a 
fleshy prolongation on each side of the tenth abdominal segment. This 
family shows a somewhat higher degree of specialization in the pro- 
thorax and labial palpi than most of the Gracilariidae. It is, however, 
not as much specialized as the species of Lithocolletis which have de- 



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veloped a cremaster, but is more like Cameraria. It may have been 
developed from the same stem as Cameraria, but its development is 
more likely to have been parallel with that of the family Gracilariidae. 
The body is considerably more chitinized, however, than in any mem- 
ber of that family. This family includes a single genus, Phyllocnistis 
Zeller, in which the pupae are from 3-4 mm. in length. 

The following species were examined: 
Phyllocnistis ampclopsisella Chambers, itisignis Frey and Boll. 

Specialized pupae with pilif en 

There are two superfamilies of Lepidoptera, the Pyralidoidea and 
the Papilionoidea, in which the pilif ers are enormously developed, and 
their presence is indicated in the pupa by lobes which extend from the 
caudo-lateral angles of the labrum towards the meson and in many 
instances are adjacent on the meson (Figs. 70, 72, 74, 76, 79; pf). 
Besides the presence of these lobes there are many other points of 
resemblance which would seem to indicate that these two superfamilies 
had a common ancestor. 

SuPERFAMiLY PYRALIDOIDEA 

This superfamily includes all those pupae which possess lobes in- 
dicating the presence of well-developed pilif ers and which do not pos- 
sess clubbed antennae. This comprises the family Pterophoridae, the 
family Attevidae, previously included in the Yponomeutidae, and 
probably all of the subfamilies of Pyralididae, although pupae of 
only six of these were examined. The Gallerinae do not possess the 
lobes indicating the presence of pilif ers, and differ in many other 
respects from most other pyralids. 

The antennae are long, at least five sixths the length of the wings, 
and in some instances extend beyond them. The maxillae and meso- 
thoracic legs are both long and extend to the caudal margin of the 
wings in most genera. The femora of the prothoracic legs are visible 
except in some genera of Pterophoridae. In all of these families the 
appendages are soldered to each other and to the body wall, but in the 
Pterophoridae they are very slightly soldered and separate readily. 
The seventh abdominal segment is free in the males of Pterophoridae 
and Attevidae but fixed in the female. In the Pyralididae it is fixed 
in both sexes. The families of Pyralidoidea may be separated as 
follows : 

a. Maxillary palpi never present ; the prothoracic and mesothoracic legs 
always extending eephalad between the sculptured eye-piece and the 
antennae; body always roughened with short spines or with small 



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70 

groups of long barbed spines and setae arising from small eleva- 
tions; dorsum of abdomen never with a deep furrow between the 

ninth and tenth segments PTEROPHORroAE. 

aa. Maxillary palpi usually present, if not, then the dorsum of the abdo- 
men with a deep furrow between the ninth and tenth segments; 
body surface seldom roughened with spines or setae, 
b. Epicranial suture never present; fronto'-clypeal suture visible for 
about half the distance between the proximal end of the antennae 
and the meson ; seventh abdominal segment free in the male and 
fixed in the female; dorsum of abdomen never with a furrow 

between segments nine and ten Attevidae. 

bb. Epicranial suture usually present, if not, then the dorsum of the 
abdomen with a deep furrow between the ninth and tenth seg- 
ments ; f ronto-clypeal suture never indicated ; seventh abdominal 
segment fixed in both sexes Pyralididae. 

Family Pterophoridae 

This family possesses a curious combination of generalized and 
specialized characters which make its position rather difficult to deter- 
mine. It has lost the maxillary palpi, the femora of the prothoracic 
legs are seldom visible, and the epicranial suture is present in but one 
genus, Pterophorus, where only a small portion of it is visible. On 
the other hand the seventh abdominal segment is free in the male and 
fixed in the female. This is clearly seen at dehiscence, for none of 
the abdominal segments possess much power of motion. The 
appendages (Fig. 70) are only slightly soldered to each other and to 
the body wall, and generally separate very readily. The wings are 
slender and pointed and, together with the other appendages, project 
slightly beyond the margin of the fourth abdominal segment. The 
clypeus, labrum, and sculptured eye-piece each bear two prominent 
setae in Pterophorus and Oxyptilus but in Platyptilia they are very 
small. There is usually a seta near the caudal margin of each gena. 
The proximal portion of each antenna is usually considerably widened 
and ridged and in Pterophorus and Oxyptilus bears long spines. The 
prothoracic legs are exceptionally long in this family and reach nearly 
to the caudal margin of the wings. The maxillae are often overlaid 
by the prothoracic legs for a part of their length, and sometimes are 
only visible for a short distance at their proximal and distal ends, the 
entire mesal portion being concealed. The location of the genital 
openings is unusual, appearing to be always on the tenth abdominal 
segment, which extends very far cephalad and forms a sort of ventral 
plate on the fixed caudal segments. In Platyptilia the plate is not so 
prominent and the dividing sutures between the segments may be dis- 



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71 

tinguished. At the cephalic margin of this plate is a large group of 
hooJced setae in Pterophorus and Oxyptilus, and in Platyptilia a 
rounded tubercle bearing four hooked setae. The abdominal spiracles 
are slightly produced. The mesothoracic spiracle is situated mesad 
of its usual position. It is also slightly produced. The peculiar spiny 
armature of most of the genera makes them very easy to distinguish 
from all other pupae. They are always found exposed, attached by 
the cremaster, and vary in length from 8-15 mm. The genera may be 
separated thus : 

a. Body with long, prominent barbed spines and setae arising mostly 
from dorsal and lateral elevations; tenth segment with a mass of 
hooked setae at its cephalic margin, 
b. Femora of the prothoracic legs exposed ; dorsal and lateral eleva- 
tions with barbed spines of varying lengths. 

Pterophorus Geoflfroy. 

bb. Femora of the prothoracic legs never exposed ; dorsal and lateral 

elevations usually with two barbed spines which are very broad 

at base and on the side of each is inserted a stout straight seta. 

Oxyptilus Zeller. 
aa. Body without any long barbed spines or setae, but with short, widely 
separated triangular projections on most of the abdominal seg- 
ments ; tenth segment with a rounded prominence near the cephalic 
margin bearing about four hooked setae Platyptilia Biibner. 

The following species were examined : 
Pterophorus paleaceus Zeller 
Oxyptilus temddactylus Fitch 
Platyptilia carduidactyla Riley 

Family Attevidae 

The genus Atteva (Figs. 72, 73), formerly included in the family 
Yponomeutidae, was found to differ in all its important characters 
from the members of that family and to be closely allied to the Pyralid- 
idae. It retains the same arrangement of setae on the clypeus and 
labrum as that in the Yponomeutidae. The setae at the caudal end of 
the body are also similar in arrangement to those in the Yponomeu- 
tidae, but the subfamily Phycitinae also have setae arranged in this 
way. It seems very probable that the Attevidae and Yponomeutidae 
arose from a common stock, but that the former branched off before 
motion was lost in the seventh segment of the male. In the Attevidae 
there is a narrow conjunctiva between the seventh and eighth segments 
in the male and there is slight motion possible. The eighth, ninth, and 
tenth segments are unusually long and distinctly segmented. There is 



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72 

no epicranial suture present, and at dehiscence the eye-pieces are not 
separated from the other face-parts, which indicates a high degree of 
specialization. The maxillary palpi are present, but not as well de- 
veloped as in most pyralids. The labial palpi are represented by a 
small polygonal area caudad of the lobes indicating the presence of 
pilifers which meet on the meson. The fronto-clypeal suture is pres- 
ent for about half the distance between the proximal ends of the an- 
tennae and the meson and it dehisces for this distance at the emergence 
of the imago. This family includes the single genus Atteva. The 
pupae of this family are from 15-20 mm. in length. 

The following species was examined : 
Atteva aurea Fitch. 

Family Pyralididae 

This family (Figs. 74, 75, 76) includes a number of subfamilies, 
of which only six are discussed here. The epicranial suture is present 
in all of these except the Epipaschiinae and a few genera of Phycitinae 
but the vertex is very short in all of the others, and often represented 
by a small triangular area, adjacent to each antenna, which does not 
reach to the meson. The antennae are long, at least seven eighths the 
length of the wings and often much longer, and the distal ends never 
meet on the meson. The labial palpi are visible only as small triangu- 
lar or polygonal areas except in the Crambinae, which often show a 
large portion between the halves of the maxillae. The maxillae are 
always long except in the Gallerinae, usually reaching the caudal mar- 
gin of the wings and sometimes extending beyond them. The maxil- 
lary palpi are present in all subfamilies except the Epipaschiinae. 
Each prothoracic leg is from one half to three fourths the length of 
the wings and its femur is always exposed. The mesothoracic legs 
generally extend to the caudal margin of the wings. The abdominal 
segments never possess spines except in the Gallerinae but are smooth 
or punctate. The spiracles are of different types, some being slightly 
produced. The location of the mesothoracic spiracle is difficult to 
determine in most specres, there being no visible opening. The ap- 
pendages are always firmly soldered to each other and the body wall. 
The pupae vary in length from 8-20 mm. The following table will 
serve to separate the subfamilies of Pyralididae : 

a. Maxillary palpi always present; epicranial suture usually distinct, 

at least for a part of its length. 

b. Maxillae never more than three fifths the length of the wings; 

dorsum of thorax and abdomen with a prominent median ridge 

and the segments covered with small spines Gallerinae. 



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73 

bb. Maxillae always more than three fifths the length of the wings ; 
dorsum of thorax and abdomen never with a median ridge or 
with small spines on the segments, 
c. Cremaster absent or never long and well developed ; furrows 
usually present on the dorsum between the ninth and tenth 
abdominal segments, or on the lateral part of the tenth seg- 
ment; head usually rounded; body never \vith a shouldered 
appearance ; labrum in its normal position, 
d. Caudal end of body with all the setae straight and very 
short; cremaster short and blunt; lateral margins of the 
dorsum of tenth abdominal segment with prominent deep 
furrows extending caudad to the proximal end of the 
cremaster; a large portion of the labial palpi often ex- 
posed Crambinae. 

dd. Caudal end of body never with all the setae straight, but 
usually long and hooked ; lateral margins of the tenth ab- 
dominal segment never with deep furrows unless they are 
extensions of the dorsal furrow between the ninth and 
tenth segments ; labial palpi never with more than a very 
small triangular or polygonal area exposed, 
e. Dorsal furrow, if present between the ninth and tenth 
abdominal segments, with a crenulate margin; meso- 
thoracic spiracles never tubular, but slit-like and not 
plainly indicated; caudal end of body without a cre- 
master, and bearing a transverse row of six or eight 

slender hooked setae Pyraunae. 

ee. Dorsal furrow usually present between the ninth and 
tenth abdominal segments but never with a crenulate 
margin, if the dorsal furrow is absent then the meso- 
thoracic spiracles produced and tubular; cremaster 

wanting or very short PHYcrriNAE. 

cc. Cremaster always present and well developed ; dorsal furrows 
never present between the ninth and tenth abdominal seg- 
ments or on the lateral part of the tenth segment ; head blunt ; 
body with a distinctly shouldered appearance ; labrum always 

cephalad of its normal position Pyraustinae. 

aa. Maxillary palpi never present ; epicranial suture never visible for any 
part of its length ; dorsal furrow between the ninth and tenth ab- 
dominal segments strongly curved caudad and apparently lined 
with a fringe of short setae EpiPASCHnNAE. 

Subfamily Gallerinae 

The pupae of this subfamily are very different from most pyralids 
and there is some doubt as to whether they should be included with 



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this family. The lobes which indicate the presence of pilifers are not 
well developed and the maxillae are short (Fig. 69). The body is 
short and thick and covered on the dorsum of. the thorax and abdo- 
men with short spines. A strongly elevated median ridge is also 
present on the thorax and on the first eight abdominal segments. The 
prothorax is very long, at least half the length of the mesothorax. 

The following species was examined : 
Galleria melonella Linnaeus. 

Subfamily Cramhinae 

The pupae of the Crambinae are easily recognized by the peculiar 
form of the short, blunt cremaster and the deep lateral grooves on the 
tenth abdominal segment. Some of the species show a large portion 
of the labial palpi, indicating that this subfamily is one of the more 
generalized. The maxillae reach almost to the caudal margin of the 
wings and the tips of the mesothoracic legs meet on the meson just 
caudad of the maxillae. The segments are almost smooth, never 
punctate. 

The following species were examined : 
Cramhtis vulgivagellus Clemens, trisectus Walker, caliginosellus Clem- 
ens. 

Subfamily Pyralinae 

The species of this subfamily resemble closely the genera Plodia 
and Ephestia of the Phycitinae. There scarcely seems to be more than 
generic differences between them. The epicranial suture is present 
and the vertex always extends to the meson. The maxillary palpi are 
well developed and usually reach the proximo-lateral angles of the 
maxillae. There is never a cremaster present, but a transverse row of 
hooked setae at the caudal end of the body. The two genera studied 
may be separated as follows : 

a. Dorsum of abdomen with a furrow between the ninth and tenth seg- 
ments, the caudal margin of the furrow distinctly crenulate. 

Pyralis Linnaeus. 

aa. Dorsum of abdomen without a furrow between the ninth and tenth 

segments. Hypsopygia Hiibner. 

The following species were examined: 
Pyralis farinalis Linnaeus 
Hypsopygia costalis Fabricius 



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Subfamily Phycitinae 

Thjs group is, for the most part, easily distinguished from other 
pyralids by the presence of the suture on the dorsum of the abdomen 
between the ninth and tenth segments, the presence of maxillary palpi, 
and, usually, of the epicranial suture. Of the genera examined, 
Ephestia and Plodia alone were without this dorsal furrow, and they 
possess tubular spiracles on the mesothorax. These two genera seem 
rather more closely related in many respects to the Pyralinae than to 
the Phycitinae. The maxillary palpi always extend to the proximo- 
lateral angles of the maxillae, and the epicranial suture is present in 
all genera examined but Pinipestis and Mineola, though it is very near 
to the suture between the head and prothorax. The vertex is usually 
represented by a small triangular area adjacent to each antenna. The 
lobes enclosing the pilifers meet on the meson in some genera. The 
genera of Phycitinae may be separated as follows : 

a. Dorsal surface without a prominent furrow separating the ninth and 
tenth abdominal segments ; mesothoracie spiracles tubular, 
b. Abdominal segments punctate ; maxillae reaching the caudal mar- 
gin of the wings Ephestia Guen6e. 

bb. Abdominal segments smooth ; maxillae never reaching the caudal 

margin of the wings Plodia Guenee. 

aa. Dorsal surface with a prominent furrow separating the ninth and 
tenth abdominal segments ; mesothoracie spiracles never tubular, 
their exact position usually difficult to determine, 
b. Body depressed ; tenth abdominal segment with the caudal end dis- 
tinctly margined and with six straight setae inserted on the ven- 
tral side of the margin Acrobasis Zeller. 

bb. Body never depressed ; tenth abdominal segment never with the 
caudal end distinctly margined or with setae inserted on the 
ventral side of the margin, 
c. Caudal end of body with four long hooked setae, and on each 
side of these a short spine or hooked seta extending laterad. 
d. Tenth abdominal segment with lateral spines very different 
from the caudal setae, 
e. Ninth abdominal segment with a lateral spine on each 
side similar to those on the tenth segment, and two 
hooked setae on the dorsum adjacent to the caudal mar- 
gin; caudal hooked setae equidistant. . , Mineola Hulst. 
ee. Ninth abdominal segment without lateral spines or 
hooked setae on the dorsum adjacent to the caudal 
margin. 

f. Caudal spines not adjacent; equidistant; of equal 
length Meroptera Grote. 



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flp. Caudal spines adjacent; two of them shorter than the 

other two Psorosina Dyar. 

dd. Tenth abdominal segment with lateral hooked setae similar 

to the caudal setae Canarsia Hulst. 

cc. Caudal end of body with a transverse row of six long hooked 
setae of equal length ; epicranial suture never present ; head 
with a prominent pointed cephalic projection. 

Pinipestis Grote. 

The following species were examined : 
Plodia interpunctella Hiibner 
Bphestia kuehniella Zeller 
Acrobasis rubrifasciella Packard 
Mineola indiginella Zeller 
Meroptera pravella Grote 
Psorosina hammondi Riley 
Canarsia ulmiarrosorella Clemens 
Pinipestis zimmermani Grote 

Subfamily Pyraustinae 

This group is distinguished by the peculiar "shouldered" appear- 
ance of the body, caused by the great width of the thorax as compared 
with the head and by the position of the labrum, which is always 
cephalad of its normal position and often located near the cephalic end 
of the body. There is never a furrow on the dorsum between the 
ninth and tenth abdominal segments. The maxillary palpi are always 
present and only a very small portion of the labial palpi is exposed. 
The epicranial suture is present in all genera. The mesothoracic legs 
and antennae, together with the metathoracic legs which are hidden 
by them, usually extend beyond the caudal margin of the wings. The 
mesothoracic spiracles often have peculiar ridges along their caudal 
margin which are sometimes covered with setae. Similar ridges are 
found in certain families of Papilionoidea and Notodontoidea. The 
shape of body and arrangement of parts in the Pyraustinae resembles 
that of certain Sphingidae, and would seem to indicate that the 
Pyraustinae are not as closely related to the Phycitinae as the other 
subfamilies, which all show a very close relationship. The genus 
Tyrausta as understood at present probably does not represent a 
natural group. Of the species studied P, fissalis and P. illibalis have 
long narrow cremasters of similar; type, while P, futilalis and P, in- 
sequalis have short broad cremasters of rather different types. There 
is also great variation in the length of the appendages, but this is not 
such a decided generic character as the form of the cremaster. This 



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subfamily includes the largest pyralids examined. The genera of 
Pyraustinae may be separated by the following table : 

a. Setae of the cremaster always hooked and equal in length to the cre- 
master or sometimes longer ; the other appendages never extending 
beyond the caudal margin of the wings, 
b. Setae of the thorax and abdomen very long, heavily chitinized, 
and forked at the distal end, usually much longer than the seg- 
ments; mesothorax and metathorax having a deep oblong pit 
with strongly chitinized edges at the base of each wing. 

PTdyctaenia Hiibner. 

bb. Setae of the thorax and abdomen never prominent, scarcely ever 

visible ; mesothorax and metathorax never having a deep oblong 

pit at the base of each wing. 

c. Mesothoracic spiracle with a prominent elevation adjacent to 

the caudal margin, which bears several tidges fringed with 

setae ; front with a distinct tubercle or small ridge at the base 

of each antenna Desmia Westwood. 

cc. Mesothoracic spiracle without any prominent elevation ad- 
jacent to the caudal margin ; front without a tubercle or ridge 

at the base of each antenna Pantagrapha Lederer. 

aa. Setae of the cremaster either straight and equal in length to the 

cremaster, or hooked and much shorter than the cremaster ; the other 

appendages often extending beyond the caudal margin of the wings. 

b. Prothorax with a distinct tubercle on each side of the meson ; cre- 

mastral setae straight and spread out fan-like. 

Tholeria Hiibner. 
bb. Prothorax without a distinct tubercle on each side of the meson ; 
cremastral setae hooked, and not spread out fan-like. 

Pyrausta Schrank. 
The following species were examined : 
Phlycta^nia ferrugalis Hiibner 
Desmia funeralis Hiibner 
Pantagrapha limata Grote and Robinson 
Tholeria reversalis Guenee 

Pyrausta fissalis Grote, illibalis Hiibner, futilalis Lederer, insequalis 
Guenee 

Subfamily Bpipaschiinae 

Only one species of this group has been examined, so no very 
definite statements can be made regarding it. The species examined 
seems to differ mainly in the absence of the maxillary palpi, which are 
present in all of the other subfamilies. The epicranial suture is not 
visible and the labrum is slightly cephalad of its normal position. The 
dorsum of the abdomen shows a decided furrow between the ninth 



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and tenth segments which is strongly curved caudad and apparently 
covered with some fine whitish setae, but this appearance may be due to 
the fine striations present. The caudal margin of the furrow is crenu- 
late. There is a very short cremaster present, bearing a small group 
of hooked setae, which are more than half as long as the tenth 
segment. 

The following species was examined : 
Lanthape platanella Clemens. 

SUPERFAMILY PAPILIONOIDEA 

The members of this superfamily are distinguished by the pos- 
session of lobes indicating the presence of well-developed pilifers and 
by their distinctly clubbed antennae. The genus Oeneis is an excep- 
tion, however, in not having the lobes well developed; but this is 
probably due to specialization, as it seems very closely allied to the 
Satyrinae, especially in the length of the prothoracic legs. Many of 
the Papilionoidea have prominent ridges and tubercles on the surface 
of the body, but there are also many genera in which the body surface 
is quite smooth and destitute of tubercles and ridges. The epicranial 
suture is present in three families, Megathymidae, Hesperiidae, and 
Lycaenidae. There has been a great deal of discussion and disagree- 
ment over the arrangement and subdivision of the families of the 
Papilionoidea. Some have divided it into two superfamilies, Hes- 
perioidea and Papilionoidea, but the pupae show no characters to war- 
rant such a division. The family Lycaenidae has been considered by 
many as the most specialized, or among the most specialized, of the 
families, yet it still retains the epicranial suture. In this family, how- 
ever, the labial palpi are entirely concealed except in the case of the 
aberrant genus Feniseca, and the shortening of the prothoracic legs is 
similar to the condition found in the Nymphalidae. It is impossible 
without further study of existing forms and a larger series of species 
to discuss fully the relationships between the different families. It is 
sufficient for the present to state that the Lycaenidae seem more nearly 
related to the generalized Hesperiidae, but have developed in a similar 
manner to the Nymphalidae, and that the Pieridae, Papilionidae, and 
Nymphalidae seem very closely related. The families of Papilionoidea 
may be separated as follows : 

a. Proximo-lateral angles of the maxillae extending laterad to the eye- 
pieces, 
b. Maxillae never reaching the caudal margin of the wings; wings 
adjacent on the meson caudad of the maxillae. . MEGATHYMroAE. 



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bb. Maxillae always extending to the caudal margin of the wings and 
sometimes beyond ; wings never adjacent on the meson. 

Hesperhdae. 
aa. Proximo-lateral angles of the maxillae never extending laterad to the 
eye-pieces, 
b. Mesothoracic legs never extending cephalad to the eye-pieces, 
c. Epicranial suture always present; head without projections; 
exposed part of maxillae never as long as the wings. 

LYCAENroAE. 

cc. Epicranial suture never present; head always with prominent 

projections; exposed part of maxillae usually as long as the 

wings. 

d. Head with two prominent projections, one at each cephalo- 

lateral angle ; metathoracic wings visible in ventral view. 

Papiuontoae. 
dd. Head with a median projection ; metathoracic wings not 

visible in ventral view Piertoae. 

bb. Mesothoracic legs extending cephalad to the eye-pieces and for a 
short distance between the sculptured eye-piece and the antenna. 

Nymphaudae. 

Family MEGATHYMroAE 

The Megathymidae, or giant skippers, are evidently the most gen- 
eralized of the Papilionoidea although they differ but little from the 
more generalized Hesperiidae, some doubt existing as to whether they 
show enough difference to warrant their being considered as a dis- 
tinct family. However, as only one pupa of this family has been seen 
by the writer and as that had lost some of the face-parts so that a 
complete description can not be given, no very definite stand can be 
taken here as to its position in the superfamily. The members of the 
superfamily Papilionoidea, as a general rule, possess but little free- 
dom of motion in the free segments and these are rarely capable of 
being telescoped. In the Megathymidae not only are the free seg- 
ments capable of a great deal of motion and of being telescoped, but 
there appears 'to be dorsal motion possible between the third and 
fourth abdominal segments, and the seventh abdominal segment ap- 
pears to possess freedom of motion in the male. The abdominal seg- 
ments are of nearly equal length, and the eighth, ninth, and tenth are 
distinctly segmented. These characters, however, appear to be re- 
tained in such generalized Hesperiidae as the genera Calpodes and 
Amblyscirtes, in which if all the above-mentioned segments do not re- 
tain freedom of motion they have certainly but recently lost it. In 
placing this family in the Papilionoidea it has been assumed that the 
pupae possess lobes indicating the presence of pilifers, but these parts 



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80 

were absent in the pupa examined. The labial palpi are represented 
by a small triangular area, but it is not known whether or not maxil- 
lary palpi are retained. The maxillae are much shorter than in the Hes- 
periidae, being only about two thirds the length of the wings, but this 
indicates nothing as to their position, as both generalized and special- 
ized pupae possess short maxillae. None of the other appendages are 
longer than the maxillae except the wings, which lie adjacent on the 
meson caudad of the maxillae. The epicranial suture is present and 
the vertex is of equal length throughout, being about one fifth the 
length of the prothorax measured on the meson. The entire body sur- 
face is covered with a whitish bloom, and on the dorsum of abdominal 
segments 7-10 there is in addition a dense covering of rather coarse 
setae. The pupa examined was 40 mm. in length and about 10 mm. 
in breadth, and belonged to the genus Megathymus. 

The following species was examined : 
Megathymus yuccae Boisduval and Le Conte. 

Family Hesperiidae 

The Hesperiidae retain considerable freedom of motion of the ab- 
dominal segments, and in many genera it would seem that dorsal 
motion is possible between the third and fourth abdominal segments 
and that the seventh abdominal segment is free in the male, or at least 
that they have only recently lost the power of motion. The epicranial 
suture is present in all genera, and the vertex is about one fifth the 
length of the prothorax measured on the meson, while the lateral mar- 
gins are considerably longer. The labrum in most genera is consid- 
erably cephalad of its normal position. The antennae never reach to 
the caudal margin of the wings but are from two thirds to three 
fourths of their length. The prothoracic legs are about half the 
length of the wings, the mesothoracic usually two thirds of the same, 
while the metathoracic pair is seldom visible. The maxillae always 
extend to the caudal margins of the wings and frequently consider- 
ably beyond. The mesothoracic spiracles usually have a peculiar kind 
of plug or plate which seems to form an external closing apparatus 
or guard, while some have prominent tubercles caudad of the opening, 
usually with a dense covering of setae. The thorax and abdomen 
usually have a more or less dense covering of setae, and some of the 
species have the entire body covered with a whitish bloom, which is 
of comparatively rare occurrence among lepidopterous pupae. The 
abdomen frequently has a furrow on the dorsum between the ninth 
and tenth segments, similar to the furrows found in the Pyralididae 
but never so deep. The cremaster in all genera is more or less trian- 



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gular, with hooked setae on the distal end, and frequently has an im- 
pressed triangular area on the dorsum. The classification of the Hes- 
periidae has long been in dispute, and with the limited amount of 
material available for examination it is impossible to state just how 
a classification of the pupae would agree with any of the proposed 
schemes. It is believed, however, that Scudder's arrangement would 
probably be followed, as the material available falls readily into his 
groups. As to the relationship between these groups there might be 
some difference of opinion. The pupae at first sight are readily divid- 
ed into two groups, one with the abdominal segments caudad of the 
fourth considerably shortened, possessing narrow flanged plates on 
the movable segments which prevent the telescoping of the body, and 
with the segmentation indistinct between the fixed caudal segments 
(Fig. yy^. This group also has the body prominently convex on the 
dorsum of the mesothorax and on the entire ventral surface of the 
thorax and abdomen. The labrum is cephalic in position. The other 
group possesses abdominal segments of more nearly equal length, hav- 
ing distinct sutures between the fixed caudal segments and the mov- 
able segments capable of being telescoped. This group has apparently 
just recently lost the power of motion in the seventh abdominal seg- 
ment of the male and dorsal motion between the third and fourth 
abdominal segments. The body is shaped like the majority of lepidop- 
terous pupae, and the labrum never quite reaches the cephalic margin 
of the body. Of this group, the genera possessing maxillae extending 
beyond the caudal margin of the wings, Calpodes (Fig. 78) and Am- 
blyscirtes, are undoubtedly more generalized, not on account of the 
maxillae, but because in all the other members of the group there is 
considerably more consolidation of the caudal abdominal segments, 
so that they seem intermediate in position between the genera men- 
tioned above and the first group. The following table will serve to 
separate the genera of Hesperiidae : 

a. Abdominal segments 5-7 never with an elevated ridge or flanged 
plate along the cephalic margin and always capable of being tde- 
scoped ; body never with a prominent convexity on the ventrid sur- 
face in the region of abdominal segments 1-4. 
,b. Maxillae extending free for a considerable distance beyond the 
caudal margin of the wings, 
c. Maxillae extending beyond the caudal margin of the body ; head 

with a long cephalic projection Calpodes Hiibner. 

cc. Maxillae never extending beyond the caudal margin of the body ; 
head never with a long cephalic projection. 

Amblyscirtes Scudder. 



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82 

bb. Maxillae never extending free beyond the caudal margin of the 
wings, 
e. Body with a dense covering of long setae and whitish bloom; 
mesothoracic spiracle with a strongly elevated oval area adja- 
cent to its caudal margin, this area chitinized in the center and 
surrounded by a dense band of short setae with a longitudinally 
striate chitinized rim forming an outer margin. 

PJiolisora Scudder. 
cc. Body sparsely covered with short inconspicuous setae ; dense 
whitish bloom never present ; mesothoracic spiracle with a some- 
what circular elevation adjacent to its caudal margin, which is 

entirely covered with setae Thanaos Boisduval. 

aa. Abdominal segments 5-7 with an elevated ridge or flanged plate 
along the cephalic margin which prevents their being telescoped ; 
body with a prominent convexity on the ventral surface in the 
region of abdominal segments 1-4. 
b. Mandibular area with distinct tubercles which are usually black 
and bearing stout setae; head slightly narrower than the meso- 
thorax. 
c. Mesothoracic spiracle semicircular in outline, the opening circu- 
lar, a broad thick band of setae around the caudal half, 
d. Dorsal furrow or depression on the ninth abdominal segment 
with its caudal margin distinctly crenulate; ventral surface 
of cremaster with an elongate furrow broadened out at the 

distal end of the cremaster Thorybes Scudder. 

dd. Dorsal furrow or depression on the ninth abdominal segment 
not distinctly crenulate ; ventral surface of cremaster with 
a triangular depression broad at the proximal end and nar- 
rowed to the distal end of the cremaster. 

Epargyrexis Hiibner. 
cc. Mesothoracic spiracle semicircular in outline, the opening cir- 
cular and surrounded by a broad thick band of setae, and 
caudad of this a distinctly elevated chitinized ridge forming an 

outer margin Cocceius Godman and Salvin. 

bb. Mandibular area smooth, without tubercles ; head as broad as the 
mesothorax Eudamus Swainson. 

The following species were examined : 
Calpodes ethlius Cramer 
Amhlyscirtes vialis Edwards 
Pholisora catullus Fabricius 

Thanaos hrizo Boisduval and Le Conte, lucilius Lintner 
Thorybes daunus Cramer 
Bpargyreus tityrus Fabricius 
Cocceiiis pylades Scudder 
Budamus pro teas Linnaeus 



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Panuly Lycaenidae 

The Lycaenidae are small pupae, between 8 and 15 mm. in length, 
which have the general shape of arctians although they are generally 
less curved on the ventral surface (Fig. 79). They retain a small 
portion of the vertex on each side and the epicranial suture usually 
touches the caudal margin of the head at the meson, making each half 
of the vertex triangular. The lobes indicating the presence of pilifers 
always meet on the meson except in the genus Feniseca. The antennae 
always extend to the caudal margin of the wings and lie adjacent on 
the meson, concealing the distal ends of the maxillae. The prothoracic 
legs are shorter than usual, varying from two fifths to one third the 
length of the wings. The mesothoracic legs are about half the length of 
the wings and the metathoracic pair are never visible. The body is 
usually quite free from projections or elevations, Feniseca being the 
only exception known, and it bears small rounded tubercles on its 
dorsal surface. The head is limited to the ventral surface of the 
body, and the suture between it and the prothorax is located on the 
cephalic margin of the body, sometimes forming a slight ridge. The 
prothorax is longer than is usual in Papilionoidea, being about half as 
long as the mesothorax. There is little, if any, motion possible be- 
tween any of the abdominal segments, and they fit together so as to 
form a smooth surface. Even the pupal skin after dehiscence shows 
no separation of the abdominal segments. The surface of the thorax 
and abdomen is covered with a reticulation of fine elevated lines with 
small papillae at their intersections and usually in the spaces between. 
These papillae usually bear cuticular appendages, of various types, 
the most peculiar being the fungiform type of the genera Chrysopha- 
nus and Heodes. There is no cremaster present in any member of 
the family. The ventral surface of the abdomen frequently bears 
groups of small hooked setae. The genital openings are usually ob- 
scured. The anal opening is peculiar, in many forms being transverse 
instead of longitudinal. The mesothoracic spii^cles are closed by a 
plug or plate which fills up the opening and usually presents a honey- 
combedo appearance. The following table will serve to separate the 
genera of Lycaenidae: 

a. Exposed portion of maxillae never more than three fifths the length 

of the wings ; cuticular appendages of the body never fungiform. 

b. Ventral surface of the body never with hooked setae caudad of 

the anal opening; thorax and abdomen usually densely covered 

with spiculate cuticular appendages ; exposed portion of maxillae 

scarcely more than half the length of the wings. 



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c. Ventral surface of ninth abdominal segment with a group of 
hooked setae on each side of the meson. 

d. Thorax with the median line slightly elevated; raised lines 
of the reticulations very prominent ; papillae short, cylin- 
drical Incisalia Minot. 

dd. Thorax with the median line never elevated ; raised lines 
of reticulations not prominent ; papillae conical. 

Uranotes Scudder. 

cc. Ventral surface of ninth abdominal segment never with hooked 

setae on each side of the meson. 

d. Ventral surface of ninth abdominal segment with a group 

of straight cuticular appendages on each side of the meson ; 

setae on body very dense and about one fourth the length 

of the abdominal segments ; papillae conical. 

Mitura Scudder. 
dd. Ventral surface of ninth abdominal segment without cutic- 
ular appendages of any kind; setae of body not dense, 
but long, usually half the length of the segment ; papillae 

short, cylindrical Thecla Fabricius. 

bb. Ventral surface of body with hooked setae caudad of the anal 
opening; papillae most numerous in the spiracular region; 
thorax and abdomen sparsely covered with short cuticular ap- 
pendages with very minute spicules ; exposed portion of max- 
illae three fifths the length of the wings, 
c. Papillae of the spiracular region confined to an area caudad 
of the spiracle ; spiracle of the second abdominal segment not 

adjacent to the wing Cyaniris Dalman. 

cc. Papillae of the spiracular region surrounding the spiracle ; 
spiracle of the second abdominal segment adjacent to the 
wing, the distance between them less than the width of the 

spiracle Rusticus Hiibner. 

aa. Exposed portion of the maxillae more than three fifths the length of 

the wings. 

b. Body of typical lycaenid shape, never flattened at the caudal end ; 

cuticular appendages fungiform. 

c. Fungiform cuticular appendages small and inconspicuous, not 

visible with a low-power lens; color light yellowish brown, 

not spotted Chrysophanus Hiibner. 

cc. Fungiform cuticular appendages large and conspicuous, easily 
visible with a low-power lens; color dark brown with black 

spots Heodes Dalman. 

bb. Body with the caudal end flattened and curved slightly ventrad. 

Feniseca Grote. 
The following species were examined : 
Incisalia niphon Hiibner 
Uranotes melinus Hubner 



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Mitura damon Cramer 

Thecla acadica Edwards, calanus Hiibner, liparops Boisduval and Le 

Conte 
Cyaniris ladon Cramer 
Rtisticus scudderi Edwards 
Chrysophanus thoe Boisduval 
Heodes hypophleas Boisduval 
Feniseca tarquinius Fabricius 

Panuly Papiuonidae 

The pupae of this family are usually long and slender, tapering 
gradually to the pointed caudal end, which is called the cremaster 
although it seldom resembles a true cremaster, and extends very little 
beyond the anal opening. The body always has two prominent cephalic 
projections, one at each cephalo-lateral angle of the head, a less promi- 
nent lateral projection on each side the metathorax at the base of each 
wing, and a low median carinate ridge which extends along the pro- 
thorax on to the mesothorax for about half its length, where it forms 
a more or less prominent projection. From this prominence the ridge 
divides, the remainder, of its course being on the metathorax, the 
divisions sometimes extending to the abdomen to form the dorso- 
lateral abdominal ridges. There is also usually present a lateral ridge 
on each side. These four ridges are continued to the end of the body, 
and are often present on the tenth segment or on the cremaster when 
absent from the remainder of the abdomen. On the ventral surface 
there is usually a ridge on each side of the face-parts, beginning at the 
cephalic projections and extending to the proximo-lateral angles of 
the maxillae. i ■ - 

The labrum is in its normal position and the lobes which indicate 
the presence of pilifers seldom meet on the meson, but are separated 
by a small portion of the labial palpi. The epicranial suture is never 
present, and the proximal ends of the antennae approach each other 
very closely on the dorsal surface of the head. The antennae never 
extend as far caudad as the wings. The wings are usually somewhat 
pointed on the ventral surface near the meson and the metathoracic 
wing is always visible here, extending for a considerable distance 
caudad of the mesothoracic wings. The maxillae always extend to the 
caudal margin of the wings. The legs are of the length usual in lep- 
idopterous pupae with the exception of the genus Iphiclides, in which 
the prothoracic legs only are about the usual length, the mesothoracic 
legs ending before the former — a very rare occurrence in this order. 
The genital openings are located in the usual positions, those of the 



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female being confluent on the ventro-meson of the eighth and ninth 
abdominal segments. Just caudad of the genital openings, at the 
cephalic margin of the tenth abdominal segment is a small tubercle on 
each side of the meson closely appressed to the surface of the body. The 
caudal end of the body bears a mass of very short hooked setae. The 
fourth, fifth, and sixth segments are movable, although they fit closely 
together to form an even surface and are not capable of telescoping. 
At dehiscence they separate, showing deep incisions. The genera of 
Papilionidae may be separated as follows : 

a. Body surface without distinctly carinate ridges; dorsal surface of 
abdomen always with a row of small rounded tubercles on each 
side of the meson, the largest on segments 4-7, and usually a row of 
smaller tubercles on each side of the spiracles ; ventral surface with 
two distinct tubercles on each leg, a transverse row near the caudal 
margin of the mesothoracic wings, and often on the veins near the 

margin PapUio Linnaeus. 

aa. Body surface with distinctly carinate ridges, but never with small 
rounded tubercles on the abdomen, occasionally with very minute 
tubercles on the wings and other appendages, 
b. Body without prominent lateral expansions of the abdominal seg- 
ments or dorsal carinate ridges, 
c. Body with a very low dorso-mesal ridge on the thorax with a 
small mesothoracic elevation; a prominent carinate ridge at 
each lateral margin of the body and no dorso-lateral ridge; 
cephalic projections large and prominent ; body very strongly 
convex on the ventral surface in the region of the wings. 

Euphoeades Hiibner. 
cc. Body with a low dorso-mesal ridge on the thorax ending in 
a very prominent mesothoracic elevation ; a very low dorso- 
lateral and lateral ridge on each side of the abdomen ; cephalic 
projections not very large ; body never strongly convex on the 

ventral surface IpTiiclides Hiibner 

bb. Body with prominent lateral expansions of the fii*8t four abdom- 
inal segments, making this the widest part of the body ; abdom- 
inal segments with dorsal carinate ridges on each side of the 
meson, most prominent on segments 5-7, which are highest in 
the middle of each segment and curve to each margin, giving it a 
scalloped appearance in lateral view Laertias Hiibner. 

The following species were examined : 

Laertias philenor Linnaeus 

Iphiclides ajax Linnaeus 

Euphoeades troilus Linnaeus, palamedes Drury 

Papilio daunus Boisduval, eurymedon Boisduval, rutultis Boisduval, 
glaucus Linnaeus, polyxenes Fabricius, thoas Linnaeus, machaon 
Linnaeus, zolicaon Boisduval, brcvicauda Saunders 



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Family Piebidae 

The pupae of this family resemble the Papilionidae very strongly 
as to the general shape of the body and arrangement of ridges and 
projections. They are much smaller, however, and are easily recog- 
nized by the fact that they possess a single median cephalic projection 
instead of two cephalo-lateral projections as do the Papilionidae. The 
epicranial suture is never present. The labrum is usually slightly 
cephalad of its normal position and a small portion of the labial palpi 
is always exposed. The maxillae vary in length from two thirds the 
length of the wings to extending slightly beyond their caudal margin. 
The legs are of normal length. The antennae are more distinctly 
clubbed than in the Papilionidae and sometimes reach the caudal mar- 
gin of the wings. The caudal end of the body is very like that of the 
Papilionidae except that the four ridges are seldom present, and the 
hooked setae are inserted in a slight concavity. The genital openings 
are in the usual positions. On the ventral surface of the tenth abdom- 
inal segment there is a low ridge, circular in outline, which encloses 
the anal opening and terminates at its cephalic end in a small tubercle 
on each side of the meson. These tubercles are located just caudad 
of the genital openings. Similar tubercles and ridges are found in the 
Nymphalidae, but they are rather more prominent in that family. 
Tubercles without the ridges occur in the Papilionidae. 

There is very little question as to whether or not the Pieridae 
and Papilionidae are related, but which is the more specialized seems 
to be a questionable point with all workers in the group. Aside from 
the question of prominences or projections, which, after all, seems a 
matter of small'importance, there is little of fundamental difference 
between the two families excepting the length of the thoracic seg- 
ments, which are more generalized in the Papilionidae, and the ridges 
and tubercles just mentioned on the ventral surface of the Pieridae, 
which resemble the Nymphalidae. The Nymphalidae certainly seem 
to be the most specialized of the Papilionoidea, although this is an- 
other much debated question. The two families have undoubtedly 
been developed from a common ancestor and represent parallel lines 
of development. The genera of Pieridae may be separated by the 
following table : 

-a. Distance from the cephalic margin of the prothorax to the distal end 
of the cephalic projection much less than the length of the pro- 
thorax ; ventral line of body often convex but never forming a prom- 
inent angle. 

b. Thorax with a strongly carinate median ridge, highest at the mid- 
dle of the mesothorax, and forming a prominent projection ; ab- 



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domen with a lateral carinate ridge on each side, forming two 
prominent projections on the second and third segments, the 
latter more prominent ; a median carinate ridge from the fourth 
abdominal segment to the caudal end of the body ; ventral line of 

body practically straight Pontia Fabricius. 

bb. Thorax without a strongly carinate median ridge, either without 
a median ridge or with one of equal height throughout, 
c. Ventral surface of body convex, but without any prominent 
rounded projection ; a low lateral ridge present along the wings, 
extending on the abdomen to the caudal end of the body. 

Eurymus Swainson. 
cc. Ventral surface of body produced into a prominent rounded 
projection which, near the caudal margin of the wings, is as 
wide as the body just caudad of the wings ; body without any 
prominent ridges, a lateral ridge present along the wings but 
scarcely indicated on any of the abdominal segments except 

the tenth Eurema Hiibner. 

aa. Distance from the cephalic margin of the prothorax to the distal end 
of the cephalic projection about equal in length to the thorax ; ven- 
tral line of body forming a prominent obtuse angle at a point about 
equidistant from the cephalic and caudal ends. .Synchloe Hubner. 

The following species were examined : 
Pontia protodice Boisduval and Le Conte, rapae Linnaeus 
Eurymus philodice Godart 
Eurema nicippe Cramer 
Synchloe genutia Fabricius 



Family Nymphaudae 

The members of this family have been variously subdivided. Some 
writers make several families of the species included here, while others 
divide them into subfamilies and tribes. At present no good charac- 
ters are known for the division of this group into families, but it must 
be admitted that the same difficulties He in the way of dividing it into 
the subfamilies and tribes proposed by Scudder ; consequently, several 
subfamily names have been introduced here to facilitate the grouping 
of the species. The Nymphalidae are distinguished from all other 
families lacking the epicranial suture by the fact that both prothoracic 
and mesothoracic legs extend cephalad to the eye-pieces and the meso- 
thoracic legs extend for a short distance between the sculptured eye- 
pieces and the antennae. The prothoracic legs are very short, rarely 
more than one third the length of the wings. The antennae and 
maxillae, except in a few instances, reach to the caudal margin of the 



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wings. The proximal ends of the antennae extend almost to the 
meson on the dorsum of the head. The labial palpi are represented 
by a very small portion caudad of the labrum and in many cases are 
entirely concealed. With the exception of Anaea andria the meta- 
thoracic wings are not visible on the ventral surface. The genital 
openings are in the usual position. The circular furrow enclosing the 
anal opening with the small tubercles caudad of the genital openings, 
is present in nearly all genera. When tubercles are present on the sur- 
face of the body they are usually on the dorsum of the abdomen and 
are arranged in seven rows, as follows: a dorso-mesal row (Fig. 8i, 
dmt), a dorso-lateral row on each side about half-way between the 
meson and the spiracles (Fig. 8i, dlt), and a dorsal (Fig. 8i, dst), 
and ventral row (Fig. 8i, vst) on each side of the abdominal spiracles. 
The subfamilies of Nymphalidae may be separated as follows : 

a. With prominent tubercles on the dorsal surface of the body, or at 
least on the abdomen, a dorso-mesal row, a dorso-lateral row on each 
side, and a row dorsad and ventrad of each row of spiracles. 

Nymphalinae. 
aa. Without prominent tubercles on the dorsal surface, at least not ar- 
ranged in rows as above, 
b. Second abdominal segment with a prominent carinate median 

elevation, somewhat constricted at its base Basilarchinae. 

bb. Second abdominal segment without a prominent, carinate median 
elevation, 
c. Body compressed, with a distinct dorso-mesal carina on the 
thorax and abdomen; ventral surface of ninth and tenth ab- 
dominal segments with hooked setae, the tubercles on the ninth 
segment covered with very short hooked setae. . .^.Apaturinae. 
cc. Body not compressed ; dorso-mesal carina never present on both 
thorax and abdomen, 
d. Abdominal segments caudad of the wings rapidly tapering and 
forming a sort of hemisphere; dorsum of abdomen with a 
prominent transverse ridge, 
e. Head with a prominent transverse ridge, extending along the 
middle of the eye-pieces and the lateral margin of the 
body; second, third, and fourth abdominal segments of 
approximately the same length; cremaster directed ven- 
trad ; transverse ridge on the fourth abdominal segment. 

Anaeinae. 
ee. Head without a transverse ridge, but with two prominent 
tubercles; second abdominal segment longer than any of 
the others; cremaster directed caudad; transverse ridge 
on the third abdominal segment tuberculate for its entire 
length EuPLOEiNAE. 



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dd. Abdominal segments caudad of the wings not rapidly taper- 
ing to form a hemisphere ; dorsum of abdomen never with a 
transverse ridge, 
e. Mesothorax prominently elevated; head with a transverse 
ridge forming slightly produced cephalo-lateral angles; 

cremaster with hooked setae Satybinae. 

ee. Mesothorax not prominently elevated ; head never with a 
tranverse ridge ; caudal end of body without hooked setae ; 
cremaster never present Oeneinae. 

Subfamily Nymphalinae 

This subfamily includes all the genera with prominent tubercles on 
the surface of the body. There are usually seven rows of these, 
mostly on the dorsal surface of the abdomen, as follows: adorso- 
mesal row ; on each side of this a dorso-lateral row ; and a row dorsad 
and ventrad of the abdominal spiracles on each side (Fig. 8i). The 
majority of species have a cephalo-lateral projection on each side of 
the head; in some these are very prominent; in others, reduced to 
small rounded tubercles or wanting. The body is usually strongly con- 
vex near the caudal margin of the wings on the ventral surface, and 
the cremaster is curved ventrad. The cremaster is more prominent 
in this subfamily than in the family Papilionidae and bears a mass of 
short hooked setae at its distal end. The species of Nymphalinae 
have been grouped into three tribes, mostly according to the size and 
arrangement of the tubercles. These three tribes may be separated 
as follows: 

a. Dorso-mesal tubercles smaller than those of the dorso-lateral rows; 
cremaster never with prominent lateral projections at the base, 
b. Cremaster longer than broad; dorso-mesal tubercles always pres- 
ent on abdominal segments 3-8 and usually on the second seg- 
ment Vanessini. 

bb. Cremaster usually broader than long ; dorso-mesal tubercles often 
wanting and never present cephalad of the fourth segment. 

Argynnini. 

aa. Dorso-mesal tubercles equal in size to those of the dorso-lateral rows ^ 

cremaster always with a lateral projection on each side at the base. 

Meutaeini. 
Tribe VANESSINI 

This tribe includes the species with all the rows of tubercles repre- 
sented and most of them complete. The tubercles of the dorso-mesal 
row are considerably smaller than those of the dorso-lateral row, 
which are usually very prominent. The rows of tubercles on either 



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side of the spiracles are always very small. The cremaster is long and 
never has prominent lateral tubercles at its proximal end. The genera 
of Vanessini may be separated by the following table : 

a. Cephalic prominences conical, with length and breadth approximately 
equal; dorso-lateral tubercles on the fourth abdominal segment al- 
ways larger than the others. 

b. Dorso-lateral tubercles on abdominal segments 2-7 long and sharp, 
spine-like, the length considerably greater than the breadth; 
dorso-mesal tubercle absent on the second abdominal segment. 

Euvanessa Scudder. 

bb. Dorso-lateral tubercles on abdominal segments 2-7 not sharp and 

spine-like ; the length scarcely, if any, greater than the breadth ; 

dorso-mesal tubercle present on the second abdominal segment. 

c. Dorso-lateral tubercles on fourth abdommal segment at least 

twice the size of the others ; median elevation of the mesothorax 

a compressed carinate ridge and usually very prominent. 

Polygonia Hiibner. 

cc. Dorso-lateral tubercles on fourth abdominal segment very little 

larger than the others; median elevation of the mesothorax 

pyramidal and not very prominent Aglais Dalman. 

aa. Cephalic prominences usually blunt, the length less than the breadth ; 

dorso-lateral tubercles on the fourth abdominal segment never 

larger than the others. 

b. Cephalic prominences broadly rounded; scarcely elevated beyond 

the outline of the body; no distinct prominence on the median 

line of the mesothorax Junonia Hiibner. 

bb. Cephalic prominences distinctly elevated beyond the outline of 
the body ; a distinct prominence on the median line of the meso- 
thorax Vanessa Fabricius. 

The following species were examined : 
Euvanessa antiopa Linnaeus 
Polygonia interrogationis Fabricius, comma Harris, faunus Edwards, 

progne Cramer 
Aglais milherti Godart 
Junonia coenia Hiibner 
Vanessa atalanta Linnaeus, huntera Fabricius, cardui Linnaeus 

Tribe ARGYNNINI 

The species included here resemble those of the preceding tribe, 
excepting that the dorso-mesal row of tubercles is only present on a 
few segments or is entirely wanting. The cremaster is short and 
broad and never has a prominent projection on each side at the prox- 
imal end. The genera of Argynnini may be separated by the following 
table : 



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a. Dorso-lateral row of tubercles of approximately equal size. 

b. Dorso-mesal row of tubercles present on abdominal segments 4-7 ; 
dorso-lateral tubercles much larger than the stigmatal rows ; cari- 

nate ridge present on mesothorax Argynnis Fabricius. 

bb. Dorso-mesal row of tubercles absent on all segments; dorso-lat- 
eral tubercles about equal in size to the dorsal stigmatal row; 

mesothorax without a carinate ridge Euptoieta Doubleday. 

aa. Dorso-lateral row of tubercles of different sizes, the largest on the 
third abdominal segment, 
b. Body with a very strong ventral curve opposite the third and 
fourth abdominal segments; mesothorax with a strongly ele- 
vated median ridge throughout its length ; head projections very 
prominent, irregularly bilobed. 

Agraulis Boisduval and Le Conte. 
bb. Body without a strong ventral curve; mesothorax with a small 
ridge on the caudal half; head projections short, pointed. 

Brenthis Hiibner. 
The following species were examined : 
/Argynnis cybele Fabricius 
Euptoieta claudia Cramer 
Agraulis vanillae Linnaeus 
Brenthis myrina Cramer 

Tribe MELITAEINI 

The species in this group have the dorso-mesal and dorso-lateral 
tubercles of approximately equal size, but none of them are very large 
and they are usually rounded. The cremaster always has a prominent 
projection on each side at its proximal end. The genera of Melitaeini 
may be separated as follows : 

a. Tubercles of the dorsal spiracular row not present on the second ab- 
dominal segment ; no tubercles present on the eighth abdominal seg- 
ment. 

b. Dorsum of abdomen with a distinct tranverse ridge on the fourth 
segment; dorsal spiracular row of tubercles not distinct on any 

of the segments Phyciodes Hiibner. 

bb. Dorsum of abdomen without a transverse ridge on the fourth seg- 
ment ; dorsal spiracular row of tubercles very large on the third 

and fourth abdominal segments Ckaridryas Scudder. 

aa. Tubercles of the dorsal spiracular row present on the second abdom- 
inal segment ; tubercles present on the eighth abdominal segment, 
b. Tubercles of the eighth abdominal segment nearly as large as the 
others, the abdominal tubercles all broadly rounded and never 
longer than broad ; cremaster with a deep depression on the dor- 
sal surface and a circular depression on the ventral surface, the 
lateral tubercles very prominent, rounded, smooth and polished. 

Euphydryas Scudder. 



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bb. Tubercles of the eighth abdominal segment much smaller than the 
others, the abdominal tubercles all somewhat pointed and longer 
than broad ; cremaster never with a deep depression on the dor- 
sal surface, but with a long narrow furrow on the ventral sur- 
face, the lateral tubercles somewhat triangular in outline and 
slightly rugose like the cremaster Cinclidia Hiibner. 

The following species were examined : 
Phyciodes tharos Drury 
Charidryas nycteis Doubleday and Hewitson 
Euphydryas phaeton Drury 
Cinclidia harrisii Scudder 

Subfamily Basilarchinae 

The genus Basilarchia differs from all the genera of Nymphalinae^ 
with which it is generally included, on account of the absence of the 
rows of tubercles. There are two cephalic projections, as in many 
Nymphalinae, and on the dorsum of the second abdominal segment 
there is a very large carinate projection. This is somewhat oval in 
outline as seen in lateral view, being constricted at the base. The body 
is not prominently excurved in the region of the appendages as in the 
Nymphalinae, but is of the same general shape. It agrees with the 
Nymphalinae only in the characters common to all members of the 
subfamily and is therefore placed in a separate subfamily. 

The following species were examined: 
Basilarclua astyanax Fabricius, arthemis Drury, archippus Cramer. 

Subfamily Apaturinae 

The species of this subfamily, included by Scudder in the Nym- 
phalinae, show no characters which unite them with that subfamily. 
The group, according to Scudder, included the genera Chlorippe and 
Anaea, which differ so widely in the pupae that they could not well be 
combined in the same subfamily. The name Apaturinae has been re- 
tained for the genus Chlorippe. These pupae are strongly compressed, 
with a prominent median dorsal carinate ridge. There are two ce- 
phalic projections and the ventral surface of the body forms a straight 
line while the dorsum is strongly arched. The antennae are slightly 
elevated and tuberculate. The genital openings are sunken and almost 
concealed. On either side of the anal opening near its cephalic end 
there is a small tubercle covered with hooked setae. The cremaster is 
short and triangular, and the hooked setae are nearly all on the ventral 
surface. 



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The following species were examined : 
Chlorippe celtis Boisduval and Le Conte, clytott Boisduval and 
Le Conte. 

Subfamily Anaeinae 

This subfamily includes a single genus, Anaea, which was included 
with Chlorippe in the tribe Apaturini by Scudder. The pupae are so 
different, however, that they have in common only the ordinary 
nymphalid characters. The body is never compressed, but the abdom- 
inal segments caudad of the wings taper very rapidly and form a 
hemisphere. The long cremaster is inserted near the center of the 
hemisphere and curves ventrad. The fourth segment has a prominent 
transverse ridge. The ventral surface of the abdominal segments 
caudad of the wings is very short and the genital openings are con- 
cealed. The head has a prominent transverse ridge at the cephalic 
end which extends caudad through the middle of the eye-pieces and 
along the lateral margin of the body. The metathorax has a rather 
prominent rounded ridge on the meson. The antennae and maxillae 
extend to the caudal margin of the wings. 

The following species was examined : 
Anaea andria Scudder. 

Subfamily Euploeinae 

This subfamily is equivalent to the family Lymnadidae of some 
authors. It includes two genera, of which only Anosia has been ex- 
amined. The general shape of the body is very like that of the genus 
Anaea of the subfamily Anaeinae, but it has the second abdominal 
segment very long, as well as the thorax, and the cremaster extends 
caudad. There is never a ridge on the head, but it has a tubercle at 
each cephalo-lateral margin. The transverse ridge is on the third seg- 
ment and is tuberculate. The maxillae do not reach the caudal margin 
of the wings in Anosia and the antennae lie adjacent on the meson 
caudad of them. 

The following species was examined : 
Anosia plexippus Linnaeus. 

Subfamily Satyrinae 

The Satyrinae are similar in shape to the Nymphalinae, but have 
no tubercles on the surface of the body and but few prominent ridges. 
The head always has a prominent transverse ridge at the cephalic end, 
and this often forms slight cephalo-lateral angles. There is also a 
slightly carinate ridge at each lateral margin of the body extending 



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as far caudad as the second abdominal segment. The metathorax al- 
ways has a median elevation, which sometimes forms a prominent 
angle. The circular ridge surrounding the anal opening is not strongly 
elevated but the tubercles are prominent on each side of the genital 
opening on the ninth segment. The genera of this subfamily may be 
separated by the following table : 

a. Cremaster broader than long, with hooked setae present on the ventral 
surface ; genital opening never with a tubercle on each side. 

Cissia Doubleday, 
aa. Cremaster longer than broad, the hooks always inserted at the dis- 
tal end, never on the ventral surface ; genital opening always with 
a distinct tubercle on each side, 
b. Mesothoracic elevation rounded; cremaster concave at tip with 
the hooked setae inserted in the hollow; body surface with fine 

indeterminate striations Cercyonis Speyer, 

bb. Mesothoracic elevation with a distinct angle ; cremaster not con- 
cave at tip ; body surface smooth Satyrodes Scudder. 

The following species were examined : 
Cissia euryttis Fabricius 
Cercyonis dope Fabricius 
Satyrodes canthus Linnaeus 

Subfamily Oeneinae 

The genus Oeneis has none of the distinguishing characters of the 
subfamily Satyrinae and therefore is not here included with the mem- 
bers of that group. The body has the general shape of a lycaenid, and 
the segments seem as devoid of motion (Fig. 80). In other respects 
it is a typical nymphalid. The antennae do not quite reach the caudal 
margin of the wings, and overlie the maxillae at their distal end, so 
that the antennae are adjacent on the meson. Their proximal ends 
are very near the meson on the dorsal surface of the head. There is 
no distinct ridge surrounding the anal opening, nor are any tubercles 
present caudad of the genital openings. There is no cremaster, nor 
hooked setae at the caudal end of the body. 

Only one species was examined : 
Oeneis semidea Say. 

Specialized pupae without pilifers 

This division includes the remaining superfamilies of Lepidoptera. 
The seventh abdominal segment is fixed in both sexes in all the fami- 
lies except the Epermeniidae, in which this segment is fixed in the 



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male. None of the species included here have dorsal movement be 
tween any of the segments cephalad of the fourth. In this they differ 
from the superfamilies Pyralidoidea and Papilionoidea, some mem- 
bers of which retain dorsal movement of the third abdominal segment. 
This group includes all the most specialized families. The origin of 
most of these is doubtful. The Noctuoidea show the strongest rela- 
tionship to the Pyralidoidea ; the Notodontoidea, to the Gelechioidea. 
All the evidence at present points to the fact that the Pyralidoidea and 
Gelechioidea have descended from a common ancestor closely allied 
to the Yponomeutoidea. The Sphingoidea and Saturnioidea, which 
show considerable relationship to each other, seem to have arisen from 
a common stem with the more generalized Bombycoidea, which in 
turn seem nearly related to the Noctuoidea and Notodontoidea. 

SuPERFAMiLY YPONOMEUTOIDEA 

The families included here show well-developed labial palpi, and 
have a large portion of the prothoracic femora exposed. All show 
the maxillary palpi except the Coleophoridae, and the same arrange- 
ment of parts prevails throughout the superfamily. The epicranial 
suture is present in all families. The prothorax is always very short 
on the meson, but much longer on each lateral margin so that each half 
is triangular. The appendages always reach beyond the caudal mar- 
gin of the fourth segment, and in some cases are almost as long as the 
body. They are soldered firmly to each other but are free from the 
body wall. Abdominal segments 1-4 are longer than any of the 
others. There are usually spines or setae present at the caudal end 
of the body but seldom a cremaster. The pupae are usually less than 
ID mm. in length. The families may be separated by the following 
table : 

a. Cremaster present, but short, with hooked setae at the distal end; 
ninth abdominal segment with a deep lateral cavity on each side; 

seventh abdominal segment free in the male EPERMENm)AE. 

aa. Cremaster absent; ninth abdominal segment never with a deep lat- 
eral cavity; seventh abdominal segment fixed in both sexes. 
b. Maxillary palpi present; caudal end of body without lateral pro- 

longations ending in spines YpoNOMEUTroAE. 

bb. Maxillary palpi never present; caudal end of body with lateral 
prolongations ending in sharp spines CoLEOPHORroAE. 

Family EPERMENm)AE 

This family, which has usually been combined with the Elachisti- 
dae, or, by some writers, with the Scythridae, is here associated with 



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the Yponomeutidae, the only important differences between the two 
families being the freedom of the seventh abdominal segment in the 
male and the presence of a very short cremaster in the Epermeniidae. 
Another difference is that in Epermeniidae the wings and other appen- 
dages are somewhat elevated at the meson and slope to each lateral 
margin. A comparison of Figure 68 with Figures 82, 83, and 85 
will show the similarity in arrangement of parts in the Yponomeutidae 
and Epermeniidae. 

The following species was examined : 
Epermenia pimpinella Murtfeldt. 

Family Ypongmeuttoae 

The genera comprising this family resemble each other very 
strongly in all important characteristics, but nevertheless possess very 
clear generic distinctions. They closely resemble certain of the gen- 
eralized gelechiids (Figs. 88, 89), and many authors have associated 
those genera with the )rponomeutids. The presence of a distinct 
fronto-clypeal suture and the peculiar arrangement of the antennae in 
the family Gelechiidae, together with the apparent loss of the labial 
palpi, seem to separate it very clearly from the Yponomeutidae, in 
which the fronto-clypeal suture is never distinct and the antennae are 
never adjacent on the meson except in Zelleria (Fig. 82). The typical 
arrangement of parts is seen in Figures 82, 83, 84, 85, 86, making 
further description unnecessary. The abdominal spiracles are all con- 
siderably produced and tubular, being longest in Plutella. There is no 
cremaster present in any of the genera. The genera of Yponomeuti- 
dae may be separated by the following table : 

a. Mesothoracic spiracles produced, tube-like; setae at caudal end of 
tenth segment hooked ; maxillae more than three fourths the length 

of the wings Plutella Schrank. 

aa. Mesothoracic spiracles not produced, slit-like; setae at caudal end 
of tenth segment straight, or occasionally slightly curved at end ; 
maxillae much less than three fourths the length of the wings, 
b. Caudal end of tenth segment showing four straight setae, generally 
two directed laterad and two caudad ; maxillary palpi touched by 
both prothoracic and mesothoracic legs, 
c. Maxillary palpi long, reaching the proximo-lateral angles of the 
maxillae ; labial palpi never becoming wider than at their prox- 
imal margin; antennae never touching on the meson. 

Yponomeuta Latreille. 

cc. Maxillary palpi short, never reaching the proximo-lateral angles 

of the maxillae ; labial palpi wider through most of their length 

than at the proximal margin ; antennae touching on the meson. 

Zelleria Stainton. 



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bb. Caudal end of tenth segment showing eight setae ; four of these on 
the ventral surface extending laterad and sometimes slightly 
curved at the tip ; mesothoracic legs never reaching cephalad to 
the maxillary palpi Argyresthia Hiibner. 

The following species were examined : 
Plutella maculipenms Curtis 
Yponomeuta padelltis Linnaeus, malinellus Zeller 
Zelleria celastrusella Kearfott 
Argyresthia freyella Walsingham 

Family Coleophoridae 

This family has usually been associated with the Elachistidae, but 
since the division of that family the name means very little unless we 
use it to include the genus Elachista and others closely related, which 
certainly would not include the Coleophoridae. They seem, rather to 
be more closely allied to the Yponomeutidae and are so considered 
here. They differ mainly in the loss of the maxillary palpi and in the 
lateral extensions of the ninth abdominal segment (Fig. 87). The ap- 
pendages are usually very long, extending nearly to the caudal mar- 
gin of the body, and often beyond it, when the movable segments are 
contracted. The abdominal segments from the first to the sixth are 
very much longer than the remaining segments. 

The following species were examined : 
Coleophora caryaefoliella Clemens, vernomaeella Chambers, maUvo- 

rella Riley. 

SuPERFAMiLY GELECHIOIDEA 

This superfamily includes those pupae which possess a distinct 
epicranial suture, with the caudal portions of the antennae lying adja- 
cent on the meson and usually separating at their distal ends to expose 
the metathoraic legs. The maxillary palpi are usually present, but 
labial palpi and prothoracic femora are seldom exposed. The body is 
usually ovate in outline as seen in dorsal or ventral view ; widest in 
the thoracic region and somewhat depressed. The superfamily is 
closely related to the* Yponomeutoidea. It includes here two groups 
representing two distinct lines of development: the group retaining 
the f ronto-clypeal suture, including the families Lavernidae, Scythri- 
dae, Gelechiidae, and Chrysopeleiidae ; and those in which it is not 
distinct or is absent, including the families Oecophoridae, Stenomidae, 
Cosmopterygidae, and Elachistidae. The latter group may represent 
a distinct superfamily when all its allied genera have been studied, but 
at present there is no evidence to warrant such a conclusion. The fol- 
lowing table will serve to separate the families of Gelechioidea : 



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a, Pronto-dypeal suture always distinct for its entire length, sometimes 
forming a prominent curve or angle at the meson, 
b. Labial palpi exposed for their entire length, 
c. Femora of the prothoracic legs visible; maxillary palpi either 
reaching the proximo-lateral angles of the maxillae or approach- 
ing them very closely; tenth abdominal segment with stout 

spines at the caudal end .LiAVERNmAE. 

cc. Femora of the prothoracic legs never visible; maxillary palpi 
minute; tenth abdominal segment with hooked setae at the 

caudal end ScYTHRmAE. 

bb. Labial palpi never exposed for their entire length, usually con- 
cealed, 
c. Maxillary palpi present and usually reaching the proximo-lateral 
angles of the maxillae ; antennae usually adjacent on the meson 
for the caudal two fifths of their length, separating at distal 

ends to show the metathoracic legs GELEcnnDAE. 

cc. Maxillary palpi never present; antennae adjacent on the meson 
for the caudal two fifths of their length but not separated at 

their distal ends Chrysopelehdae. 

aa. Pronto-clypeal suture never distinct for its entire length and never 
reaching the meson, 
b. Abdominal segments 4-6 movable, with very deep incisions be- 
tween the segments on the dorsal and ventral surfaces; body 
depressed, 
c. Maxillary palpi large, usually reaching the proximo-lateral 
angles of the maxillae; hooked setae never present on the ven- 
tral surface of the ninth abdominal segment OEcoPHORmAE. 

cc. Maxillary palpi minute ; hooked setae always present on the ven- 
tral surface of the ninth abdominal segment Stenomtoae. 

bb. Abdominal segments 4-6 never all movable; incisions between 
the segments of equal depth on all surfaces ; body not depressed, 
c. Maxillary palpi present ; sixth abdominal segment movable. 

CoSMOPTERYQroAE. 

cc. Maxillary palpi absent; no abdominal segments movable. 

Elachistidae. 

Family LAVERNmAE 

The pupae belonging to this family are more generalized than any 
other members of the superfamily and closely resemble pupae belong- 
ing to the family Yponomeutidae. They have been included here on 
account of the distinct fronto-clypeal suture, present in all except the 
more specialized Gelechioidea, and also because the prothorax, which 
is so short on the meson in Yponomeutidae with each half triangular 
in outline, in this family loses that condition and becomes almost as 
long on the meson as at the lateral margins, so that each half is sub- 



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100 

quadrangular. This is the only family of Gelechioidea which retains 
both labial palpi and exposed portions of the prothoracic femora. The 
appendages have the characteristic arrangement of the superfamily 
(Fig. 88) except that the antennae do not separate near their distal 
end to expose a portion of the metathoracic legs, but these are seen 
caudad of the antennae adjacent on the meson. The wings are long 
and pointed in Lophoptilus, but rounded in Laverna. The first four 
abdominal segments are longer than the remaining caudal segments 
in this family, and the appendages are soldered to them, but are free 
for the remainder of their length. This family has been considered as 
a subfamily of Elachistidae by most authors, and has usually included 
Cosmopteryx, which is a much more specialized genus. The following 
table will serve to separate the genera of Lavemidae : 

a. Head long, somewhat pomted, the length more than half the greatest 
width ; f ronto-clypeal suture making an acute angle at meson ; spines 
of the tenth segment extending dorsad and not visible in ventral 
view ; exposed part of metathoracic leg about one fifth the length of 
the portion of the antennae lying adjacent on the meson. 

Lophoptilus Sircom. 
aa. Head short, blunt, the length about equal to half the greatest widtli ; 
f ronto-clypeal suture making an obtuse angle at meson; spines of 
tenth segment extending laterad and visible in ventral view ; ex- 
posed part of metathoracic leg about equal in length to the portion 
of the antennae lying adjacent on the meson Laverna Curtis. 

Family Scythrtoae 

The pupae of this family also resemble the Yponomeutidae in some 
respects, and have been included with them by some authors. Others 
have associated the family with the Elachistidae, while Stainton in- 
cluded it with the Gelechiidae as the genus Butalis. The antennae in 
this family meet on the meson, but do not separate to show the distal 
ends of the metathoracic legs as is the general rule in this superfamily. 
Instead, the mesothoracic wings lie adjacent on the meson caudad of 
the antennae, and the appendages are firmly soldered to each other and 
to the body (Fig. 89). As the appendages extend caudad for about 
half the length of the seventh abdominal segment, it follows that there 
can be no motion possible between any of the abdominal segments, 
unless there be slight dorsal motion. The prothorax is typically gele- 
chiid in character. The abdominal spiracles are considerably pro- 
duced and tubular, their length varying in the different species. The 
setae of the body are nearly all hooked, and a few longer ones are pres- 



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101 

ent at the caudal end of the body. Only one genus of this family was 
available for study. 

The following species were examined : 
Scythris eboracencis Zeller, impositella Zeller. 

Family Gelechiidae 

The pupae of the Gelechiidae never show any portion of the labial 
palpi, unless it should be a very small triangular area caudad of the 
labrum, between the halves of the maxillae. The prothoracic femora 
are never exposed (Figs. 90, 91, 92, 93, 94). The fronto-clypeal su- 
ture is always distinct and usually extends almost straight across be- 
tween the proximal ends of the antennae, but occasionally each half 
is directed cephalad near the meson so that an angle is formed at their 
junction. The caudal parts of the antennae usually lie adjacent on the 
meson for about two fifths of their length and usually cover the caudal 
ends of the maxillae and sometimes of the prothoracic and mesotho- 
racic legs. They usUally separate at their distal ends to show the meta- 
thoracic legs, or what will be referred to as such in this paper. There 
was not enough available material in condition for dissection to de- 
termine whether the maxillae ever reached the caudal margin of the 
wings and overlaid the metathoracic legs, as might easily be the case. 
The maxillary palpi are present in all, but do not always reach the 
proximo-lateral angles of the maxillae. They are always reached by 
both prothoracic and mesothoracic legs. The wings vary somewhat 
in length, but are usually firmly soldered down, and the abdominal seg- 
ments are somewhat depressed on the ventral surface, forming a 
shallow cavity into which the wings are fitted and, therefore, are not 
elevated above the surface of the body. There are usually very deep 
incisions between the segments, especially on the dorsal and ventral 
surfaces. Many species have the incisions deeper on the ventral sur- 
face so that the caudal end of the body may be strongly curved ven- 
trad. The pupae are usually very active, and many of them are able 
to move after the fashion of click-beetles. The body is entirely cov- 
ered with setae in some genera, while others have a fringe of setae 
along the margin of certain slightly projecting ridges and occasional 
depressions found usually on the seventh abdominal segment. There 
seems doubt as to the generic standing of the following species : Aris- 
totelia physaliella, Gnorimoschema lavernella, and Recurvaria variella; 
at least they differ from other species examined in these genera. In 
the case of Aristotelia it has been impossible to determine which 
species is the type of the genus. The genera of Gelechiidae may be 
separated as follows : 



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102 

a. Body setae very long and heavily chitinized, often as long as the seg- 
ments of the abdomen, and, as seen under high power, usually forked 
at the apex ; f ronto-clypeal suture curving cephalad from the proxi- 
mal ends of the antennae to the cephalic margin of the body; crc- 
master always present; dorsal cephalic margin of segments two, 
three, and four of the abdomen with a slight rounded projection on 
each side of the meson, edged with a dense fringe of whitish setae di- 
rected cephalad, caudad .of this a prominent elevation bearing a sim- 
ilar fringe of setae on the summit (Fig. 91). 

b. Dorsal surface of the cephalic margin of the movable segments 
with a prominent cavity on the meson having heavily chitinized 

edges Trichotaphe Clemens. 

bb. Dorsal surface of cephalic margin of the movable segments with- 
out any cavity ; a strongly chitinized ridge separating the ce- 
phalic margin from the remainder of the segment, cephalad of 
this a band of short spines, then a prominent furrow, the furrow 
and the remainder of the cephalic margin being deeply punctate. 

Ypsolophus Fabricius. 
aa. Body setae never modified; segments two, three,* and four of the ab- 
domen never as described in a. 
b. Entire body with a dense covering of whitish setae visible to the 
unaided eye, giving the pupa a furred appearance, 
c. Maxillary palpi long, reaching the proximo-lateral angles of the 
maxillae; antennae lying adjacent on the meson for about two 
fifths of their length; cremaster short and blunt, the end set 
with about eight stout curved setae ; length 8-10 mm. 

Anacampsis Curtis. 

cc. Maxillary palpi short, never reaching the proximo-lateral angles 

of the maxillae ; antennae just meeting on the meson ; cremaster 

with a sharp point curved dorsad, with curved setae at the sides 

and base ; length 5-6 mm Aristotelia (a)Hubner. 

bb. Entire body never covered with setae, 
c. Seventh abdominal segment with a dense fringe of setae on some 
portion, 
d. Fringe of setae confined to the cephalic and lateral edges of a 
prominent lateral cavity, 
e. Body smooth, not depressed ; cephalic edge of the lateral cav- 
ity trilobed (Fig. 93) Evippe Chambers. 

ee. Body with small spines, strongly depressed; cephalic edge 

of the lateral cavity bilobed Telpiiusa Chambers. 

dd. Fringe extending around the segment or nearly so. 

e. Fringe extending around the segment in a straight line ; body 
not noticeably depressed, the surface smooth. 

Recurvcaria (a) Haworth. 
ee. Fringe extending around the segment in a more or less wavy 
line; body noticeably depressed and very broad in the 
thoracic region, surface with punctures or spines. 



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103 

f . Abdominal segments 8-10 distinctly tapering to the cau- 
dal end of the body ; fringe of seventii segment extend- 
ing in a wavy line around the body and edging two very 
large lobes on the dorsal surface. 

Trypanisma Clemens. 

flp. Abdominal segments 8-10 not tapering but blunt and 

slightly rounded at the caudal end of the body; fringe 

extending in a wavy line around the seventh segment, 

without prominent lobes on the dorsal surface. 

Gelechia Hiibner. 
cc. Seventh abdominal segment without any fringe of setae, 
d. Caudal end of body with short, stout projecting spines, 
e. Caudal end of body with one such spine on the dorso-meson, 
projecting dorsad ; f ronto-dypeal suture almost straight. 

PhtJiorimaea Meyrick. 

ee. Caudal end of body with a median spine and one on each 

lateral margin; f ronto-dypeal suture extending cephalad 

from the base of each antenna to the cephalic margin of 

the head Sitotroga Heinemann. 

dd. Caudal end of body with straight or curved setae, 
e. Hooked setae present at the caudal end of the body, 
f . Antennae reaching the caudal margin of the wings, their 
caudal ends separated to show the metathoracic legs, 
g. Caudal end of body with at least five long hooked setae 
on each side of the meson ; antennae slightly enlarged 
at their proximal ends, making the cephalic end of the 

body somewhat truncate Recurvaria (b) Haworth. 

gg. Caudal end of body never with more than two long 
curved setae on each side of the meson ; cephalic end 

of body rounded Aristotelia (b) Hiibner. 

flp. Antennae never reaching the caudal margin of the wings 
nor separating at their caudal ends to expose the meta- 
thoracic legs Gnorimoschema (a) Busck. 

ee. Hooked setae never present at the caudal end of the body ; 
a few short, straight setae present. 

Onorimoschema (b) Busck. 

The following species were examined : 
Trichotaphe flavocostella Clemens 

Ypsolophns citrifoUellus Chambers, eupatoriellus Chambers 
Anacampsis sp., rhoifructella Clemens 
Aristotelia (a) salicifungiella Clemens 
Aristotelia (b) physaliella Chambers 
Bvippe prunifoliella Chambers 
Telphusa quercimgracella Chambers, palliderosacella Chambers 



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104 

Recurvaria (a) apicitripunctella Clemens 

Recurvaria (b) variella Chambers 

Trypanisma prudens Clemens 

Gelechia cercerisella Chambers, discoocellella Chambers, serotinella 

Busck 
Phthorimaea sp. 
Sitotroga cerealella Olivier 
Gnorimo schema (a) lavernella Chambers 
Gnorimoschema (b) gallaesolidaginis Riley 

Family CHRYSOPELEaiDAE 

This family includes the genus Chrysopeleia which was formerly 
included with the Elachistidae. It has the same arrangement of parts 
as that in the genus Elachista but retains the f ronto-clypeal suture and 
has no cremaster (Fig. 95). Until more is known of the relationships 
of this genus it seems better to place it in a family by itself. The ap- 
pendages are slightly elevated and firmly soldered to each other and to 
the body. They extend well on to the seventh abdominal segment, so 
that there appears to be no motion possible between any of the body 
segments. There is no cremaster present and only a few short straight 
setae at the caudal end of the body. The abdominal spiracles are pro- 
duced and tubular. The pupae are very small, being only about 3 mm. 
in length. 

The following species was examined : 
Chrysopeleia ostryaeella Chambers. 

Family OECOPHORroAE 

This family (Figs. 97 and 98) includes those pupae in which the 
fronto-clypeal suture is not present for its entire length and which 
have large maxillary palpi, usually reaching the proximo-lateral angles 
of the maxillae. All the species examined showed the presence of very 
fine setae arranged in groups over the surface of the abdomen, but 
these were hard to locate in Psilocorsis. The incisions between the 
segments are very deep in all members of the family. Of the three 
genera studied Psilocorsis, Agonopteryx, and Depressaria, the first 
seemed more nearly related to the Stenomidae, while the other two 
were typical gelechiids, except that the fronto-clypeal suture was never 
distinct. It seems probable that a revision of the group might sepa- 
rate these genera. The table to genera will indicate the principal dif- 
ferences. The body is usually strongly depressed and 5-10 mm. in 
length. The following table will serve to separate the genera: 



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105 

a. Femora of the prothoracic legs exposed. 

b. Wings pointed, extending on to the sixth abdominal segment; 
proximal end of each antenna elevated and roughened with trans- 
verse ridges; cremaster present PsUocorsis Clemens. 

bb. Wings not pointed, and never extending beyond the caudal mar- 
gin of the fourth abdominal segment ; proximal end of each an- 
tenna never elevated or roughened with transverse ridges; cre- 
master never present Depressaria Haworth. 

aa. Prothoracic femora never exposed Agonopteryx Hiibner. 

The following species were examined : 
Psilocorsis obsoletella Zeller, quercicella Clemens 
Depressaria heracliana De Geer 
Agonopteryx nebulosa Zeller 

Family Stenomidae 

The Stenomidae include most of the genera formerly grouped 
under the name of Xylorictidae. The appendages are arranged as in 
the Gelechiidae and the maxillary palpi are minute (Fig. 96). A 
small portion of the labial palpi is usually apparent. The appendages 
are firmly soldered to each other and to the body wall. They extend 
slightly beyond the caudal margin of the fourth abdominal segment, so 
that there are three free segments, the fourth, fifth, and sixth. The 
margins of these free segments are serrate along the edges of the in- 
cisions, which are very deep, especially on the ventral surface, and per- 
mit the caudal end of the body to be very sharply curved ventrad, 
reaching almost to the caudal margin of the wings (Fig. 96a). The 
f ronto-clypeal suture is visible for a short distance mesad of the prox- 
imal end of each antenna, but it never reaches the meson. There are 
many curved setae present on the ventral surface of the ninth abdom- 
inal segment. The body is always more or less depressed, and in 
Stenoma is about 8 mm. in length, in Menesta 3 mm. The genus 
Menesta, formerly included in the Gelechiidae, seems more closely 
allied to Stenoma and is included here. Stenoma possesses peculiarly 
modified setae on the body surface. These genera may be separated as 
follows : 

a. Antennae modified at their proximal ends, forming an enlarged cor- 
rugated area ; hooked setae on the ventral part of the ninth abdom- 
inal segment never on a distinct prolongation Stenoma Zeller. 

aa. Antennae never modified at the proximal end; hooked setae on the 
ventral part of the ninth abdominal segment on a distinct prolon- 
gation Menesta Clemens. 



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106 

The following species were examined: 
Stenoma schlaegeri Zeller 
Menesta albaciliaeella Chambers 

Family Cosmopteryqidae 

This family name as used by most authors is equivalent to Laver- 
nidae or Momphidae, and the genera included under all these names 
are usually associated with each other. The Cosmopterygidae 
are much more specialized, however, as they retain neither visible 
labial palpi nor prothoracic femora (Fig. 99). The appendages are 
firmly soldered to each other and to the body wall as far as the caudal 
margin of the sixth abdominal segment, which allows freedom of 
movement to this segment. There are some generalized characters 
present, however — ^the length of the first six abdominal segments, 
which are as long as in Yponomeutidae, and the shape of the pro- 
thorax, which is shorter on the meson than at each lateral margin. The 
abdominal spiracles are slightly produced and tubular. There is a very 
short cremaster present bearing eight hooked setae, of which four are 
longer than the remainder. The pupae are about 4 mm. in length. 

The following species was examined : 
Cosmopteryx clandestinella Busck. 

Family Elachistidae 

This family has been variously subdivided in the past few years, 
for, like the Tineidae, it included a large number of species which did 
not form a natural group. Some authors do not retain this family 
name, but as the nomenclature of the ^group appears to be still in a 
rather unsettled condition, this name is retained for the present to in- 
clude the genus Elachista. The appendages are arranged as in other 
gelechiids, but there is no trace of maxillary palpi (Fig. 100). The 
surface of the body is covered with large rounded tubercles and the 
dorsal surface shows three distinct longitudinal elevations or ridges, 
one on the meson and one near each lateral margin, bearing the spir- 
acles on the summit. The wings and other appendages are firmly sol- 
dered to each other and to the body wall, and there appear to be no 
free segments. The prothorax is typically gelechiid and the meso- 
thorax shows a decided alar furrow on each side. There is a distinct 
cremaster present, but it shows no hooked setae. The pupae are sus- 
pended from a stem or leaf after the manner of some papilionids, with 
a silken girth around the body. The pupae average 3.5 mm. in length. 

The following species was examined : 
Elachista praelineata Braun. 



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107 

SuPERPAMiLY NOCTUOIDEA 

This superfamily includes three families, Noctuidae, Liparidae, 
and Arctiidae. The Syntomidae also belong to this group, but as only 
one species of this family was examined and this ^owed no charac- 
ters to separate it from the Arctiidae, the Syntomidae are not dis- 
cussed as a separate family. The Noctuoidea and Bombycoidea in- 
clude all the specialized families which retain labial palpi. The fami- 
lies of Noctuoidea may be separated thus : 

a. Body seldom with setae arranged around scars of larval vernicae, if 
present, then the femora of the prothoracic legs exposed, or a long 
cremaster present bearing hooked setae at the distal end ; prothoracic 
femora usually visible, if not, then the mesothoracic leg usually ex- 
tending cephalad to the eye-pieces Noctuidae. 

aa. Body always with setae arranged aroimd the scars of larval verrucae ; 
femora of the prothoracic legs never visible, 
b. Maxillae never more than two fifths the length of the wings ; body 

setae conspicuous; labial palpi usually visible LiPARmAE. 

bb. Maxillae two thirds the length of the wings, or longer; body setae 
inconspicuous; labial palpi seldom visible Argthdae. 

Family Noctutoae 

This family (Figs. loi, 102, 103), with a few exceptions, is char- 
acterized by the presence of labial palpi and of maxillae which extend 
to the caudal margin of the wing, or very closely approximate this 
length. Very many of the genera have a large portion of the pro- 
thoracic femora exposed. .Those which do not show any portion of 
the prothoracic femora have the mesothoracic leg extending cephalad 
to the eye-pieces, with a few exceptions in the genera Homopyralis, 
Plusiodonta, and Anomis. Those lacking labial palpi have setae ar- 
ranged around the scars of larval verrucae, as in the Arctiidae. They 
differ from the Arctiidae in having hooked setae on the cremaster, 
and in lacking flanged plates on the abdominal segments. Maxillary 
palpi are found in some members of the subfamilies Agrotinae, Cucul- 
lianae, and Hypeninae. Since there was not enough material available 
for study to furnish a basis for subfamily characters, the genera have 
been grouped as seemed best for purposes of classification. As far as 
possible the names of subfamilies as used by Hampson in the "Cata- 
logue of Lepidoptera Phalaenae" have been adopted. This arrange- 
ment could not be followed throughout, however, and so it must be re- 
membered that the subfamily names used here are adopted as a matter 
of convenience and do not stand for the genera which Hampson 



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108 

grouped together. His generic and specific names have been adopted 
as far as possible. As to the phylogeny of the group, too little ma- 
terial has been examined to warrant a decided opinion on the subject. 
It seems probable, however, that most of the subfamilies discussed rep- 
resent the ends of many lines of development. There are various 
stages of development fotmd in all groups, and there are some mem- 
bers of each subfamily studied, except the Phytometrinae and Momi- 
nae, which show the epicranial suture. 

The subfamilies mentioned above which retain maxillary palpi are 
undoubtedly the most generalized, the Mominae, which show neither 
labial palpi, prothoracic femora, maxillary palpi, nor an epicranial su- 
ture, are undoubtedly the most specialized, but nothing can be said 
with certainty as to the other groups. No attempt has been made to 
arrange the subfamilies in phylogenetic order either in the tables or in 
the discussion of subfamilies. The subfamilies of Noctuidae may be 
separated by the following table : 

a. Prothoracic legs reaching cephalad to the eye-piece, mesothoracic legs 

never reaching as far cephalad ; prothoracic femora usually visible. 

b. Cremaster, or caudal end of the body, with all the setae curved or 

hooked, never with any long straight setae. 

c. Setae of the cremaster usually all of the same size and length, 

either slightly curved or hooked; cremaster* never with six or 

eight hooked setae of which the mesal ones are larger and longer 

than the remainder Acronyctinae. 

cc. Setae of the cremaster or caudal end of the body usually of two 
sizes, 
d. Body never with scars of larval verrucae bordered with setae ; 
labial palpi and prothoracic femora always visible, 
e. Wings and maxillae produced at meson into a distinct 
rounded projection extending on to the fifth abdominal seg- 
ment; labrum located near the cephalic end of the body; 

body never heavily chitinized Phytometrinae. 

ee. Wings and maxiUae never extending beyond the caudal 
margin of the fourth segment ; labrum in the normal posi- 
tion; body always heavily chitinized Cuculuanae. 

dd. Body with rounded or oval areas bordered with setae, as in 
the Arctiidae, especially noticeable on the dorsum of the 
abdomen and in the spiracular region ; labial palpi and pro- 
thoracic femora never visible Mominae. 

bb. Cremaster or caudal end of the body never with all the setae 

curved or hooked. 

c. Cremaster or caudal end of body with at least two long straight 

stout setae, 1-2 mm. in length, rarely with additional hooked 

setae Hadeninae. 



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109 

cc. Cremaster or caudal end of body never with long straight stout 
setae, 
d. Cremaster bifurcate, narrowed at the caudal end; dorsum of 
movable abdominal segments with one or more rows of deep 

circular pits with dark, chitinized margins Agrotinae. 

dd. Cremaster short, broader at the distal end, very thin and 
plate-like, the caudo-lateral angles produced into short 
rounded lobes, with two or three small rounded projections 
between often bearing small delicate setae; dorsum of tbe 
movable abdominal segments never with pits ; body strongly 

rugose, the abdominal segments spinose Aqaristinae. 

aa. Prothoracic and mesothoracic legs both reaching cephalad to the eye- 
piece or to the maxiUary palpus where this is present ; prothoracic 
femora seldom visible, 
b. Cremaster usually present; curved or hooked setae always pres- 
ent at the caudal end of body, usually eight in number. 

c. Body always covered with a whitish bloom Catocaunae. 

cc. Body never covered with a whitish bloom Hypeninae. 

bb. Cremaster or setae never present at the caudal end of the body ; 
fifth abdominal segment with a row of spines along the caudal 
margin extending from the sculpturing of the dorsum almost to 
the meson on the ventral surface Sarrothripinae. 

Subfamily Agrotinae 

This subfamily, as here considered, includes those pupae with a 
stout, rugose, more or less bifurcate cremaster, and with a row of 
large circular pits with heavily chitinized margins along the cephalic 
margin of some of the abdominal segments, usually between the fourth 
and seventh (Fig. loi). In these pupae, the prothorax is very long, 
at least two thirds the length of the mesothorax, and the epicranial 
suture is sometimes present in the genus Agrotis. Labial palpi are al- 
ways present and exposed for their entire length, and the prothoracic 
femora are seen in some of the genera. The mesothoracic legs, an- 
tennae, and maxillae are of practically the same length and usually ex- 
tend to the caudal margin of the wings. The metathoracic legs are 
seldom visible and only the prothoracic legs extend cephalad to the eye- 
pieces, and these do not separate the sculptured eye-piece and antenna. 
The body is stout, and when retracted the length is about three times 
the width. The genera of Agrotinae may be separated as follows : 

a. Femora of the prothoracic legs visible. 

b. Dorsal cephalic margin of the movable abdominal segments with 
a single even row of pits, numbering less than fifteen ; epicranial 
suture present ; cremaster often with a row of four setae near its 
proximal end Agrotis Qphsenheimer. 



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110 

bb. Dorsal cephalic margin of the movable abdominal segments with 
an interrupted row of pits numbering more than fifteen ; epicra- 
nial suture never present; cremaster never with a row of four 

setae near its proximal end Hapalia Hiibner. 

aa. Femora of the prothoracic legs never visible Noctua Linnaeus. 

The following species were examined : 
Agrotis badinodis Grote, bicarnea Guenee 
Hapalia incivis Guenee 
Noctim clandesHna Harris 

Subfamily Cucullianae 

As only two specimens of one genus, Graptolitha, were available 
for study, little can be said as to subfamily characters. These speci- 
mens differ from members of other subfamilies except the Catocalinae 
in having all the setae at the caudal end of the body hooked. There 
are two setae at the meson very much larger and more heavily chi- 
tinized than the remaining setae, which are usually four in number. 
In other respects, as the length of prothorax, size and shape of body, 
arrangement of appendages, presence of epicranial suture and labial 
palpi, exposed femora of the prothoracic legs, and traces of maxillary 
palpi, they resemble the Agrotinae and especially the Hadeninae as 
the movable abdominal segments are finely punctate along their ce- 
phalic margin. 

The following species were studied : 
Graptolitha laticinerea Grote, antennata Walker. 

Subfamily Hadenituie 

This subfamily includes pupae having stout straight setae or spines 
at the caudal end of the body. There are usually two, from 1-2 mm. 
in length, and they may be inserted in a short cremaster or directly in 
the caudal end of the body (Fig. 102). One genus, Cirphis, has addi- 
tional slender hooked setae. The prothorax is very long, as in Agro- 
tinae, at least two thirds the length of the mesothorax. The epicra- 
nial suture is present in the genera Polia, Hadena, Lycophotia, and 
Eriopus. The appendages, which in Agrotinae are of the same length 
and generally reach the caudal margin of the wings, are in this sub- 
family unequal in length. The maxillae usually reach the caudal mar- 
gin of the wings, but the mesothoracic legs are shorter, and the anten- 
nae in some forms equal these or are very much shorter. Except for 
Cirphis and Monima the abdominal segments are punctate. These 
two genera have the movable abdominal segments pitted as in the 



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Agrotinae. The prothoracic leg extends cephalad to the eye-piece, but 
there is never a long point extending cephalad between the antennae 
and the sculptured eye-piece. The genera of Hadeninae may be sep- 
arated by the following table : 

a. Dorsum of movable abdominal segments with a row of deep pits along 
the cephalic margin. 

b. Caudal end of body with a very short cremaster ending in two 
straight sharp setae with four slender hooked setae at tiie prox- 
imal end, the hooked setae about half the length of the straight 

ones Cirphis Walker. 

bb. Cremaster very short, with a long straight seta inserted in each 
caudo-lateral angle; hooked setae never present. 

Monima Hiibner. 
aa. Dorsum of movable abdominal segments never with a row of deep 
pits along the cephalic margin, 
b. Mesothoracic leg never reaching cephalad to the eye-pieces; pro- 
thoracic femora always exposed, 
c. Caudal end of the abdomen with four closely approximated spines 

of the same size and shape Ehodophora Guenee. 

cc. Caudal end of body with only two spines, 
d. Spiracles noticeably modified, usually with a prominent de- 
pression near their caudal margin; spiracular opening di- 
rected somewhat caudad; cephalic third of the movable 
abdominal segments usually slightly elevated and densely 
punctate. 
e. Cremaster slender, pointed, spines closely approximated; 
abdominal spiracles with a darker, elevated, crescent- 
shaped area almost surrounding the spiracle and with a 
deep cavity larger than the spiracle directly caudad ojE it; 

mesothoracic spiracles not modified Meliana Curtis. 

ee. Cremaster short, blunt; area around each abdominal spir- 
acle slightly elevated and darker in color, a prominent cav- 
ity caudad of the spiracle with a chitinized ridge along the 
caudal margin of the spiracle ; mesothoracic spiracles also 
modified, the opening extending half the distance between 
the antennae and the meson, 
f . Chitinized ridge along the caudal margin of the abdominal 
spiracles distinctly serrate; clypeal region not promi- 
nently elevated and without deeply impressed lines. 

Laphygma Guen6e. 
ff. Chitinized ridge along the caudal margin of the abdominal 
spiracles not distinctly serrate; clypeal region promi- 
nently elevated and with deeply impressed lines. 

Prodenia Guen6e. 



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dd. Spirades normal, without any prominent elevations or de- 
pressions adjacent ; cephalic third of the movable abdominal 
segments usually not elevated ; segments punctate, 
e. Epicranial suture present and distinct, 
f . Cephalic third of movable abdominal segments with fine 
punctures, the remainder of the segments smooth; no 

cremaster present Lycophotia Hiibner. 

fF. Abdominal segments all finely punctate and with wavy 
impressed lines, more densely punctate along cephalic 
margin of the movable segments ; a short, distinct rugose 

cremaster present Hadena Schrank. 

ee. Epicranial suture wanting, 
f . Abdominal segments 5-7 very finely punctate on the ce- 
phalic third Chloridea Westwood. 

ff. Abdominal segments with very large, shallow punctures, 
at least on the cephalic third of segments 4-7. 
g. Abdominal segments all punctate ; segments 4-7 having 
the punctures very large on at least the cephalic two 
thirds of each segment and fine punctures on the re- 
mainder Pyrrkia Hiibner. 

gg. Abdominal segments 1-7 punctate ; segments 4-7 with 
larger punctures on the cephalic half of each segment 
and usually with finer punctures on the remainder. 

Polia Ochsenheimer. 

bb. Metathoracic leg reaching to the eye-piece; cephalic margin of 

tenth abdomincd segment with a row of deep oblong pits ; protho- 

racic femora never exposed Eriopus Treitschke. 

The following species were examined : 
Cirphis unipuncta Haworth, phragmitidicola Guenee 
Rhodophora gaurae Smith and Abbot, florida Guenee 
Meliana albilinea Hubner. 
Laphygma frugiperda Smith and Abbot 
Prodenia ornithogalli Guenee 
Lycophotia margaritosa Haworth 
Hadena vulgaris Grote 

Chloridea obsoleta Fabricius, virescens Fabricius 
Pyrrhia umbra Hiifnagel 

Polia retngera Stephens, picta Harris, meditata Grote 
Briopus floridensis Guenee 
Monima alia Guenee 

Subfamily Agaristinae 

The members of this subfamily show remarkable uniformity and it 
is rather difficult to separate the genera. They have been given family 



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113 

rank by some authors, and while the species included here differ from 
the typical noctuid in many respects, still no structural characters 
could be found to warrant their separation from the Noctuidae. The 
entire body surface is very rough and spinose, while the cremaster 
is short, broad, and decidedly flattened, with its caudo-lateral an- 
gles produced into rounded lobes, and with the caudal margin often 
crenulate. There are sometimes short straight setae present along 
the caudal margin. The antennae, mesothoracic legs, and maxillae 
usually reach the caudal margin of the wings, or approach it very 
closely. Each prothoracic leg extends cephalad to the eye-pieces. The 
epicranial suture is always present. The labial palpi are always visi- 
ble, but the femora of the prothoracic legs are concealed except in 
occasional specimens. The abdominal spiracles are somewhat elevated 
and are surrounded by a heavy, dark, chitinized border. The open- 
ings appear to be fringed with fine setae. The genera of this sub- 
family may be separated as follows : 

a. Antennae always reaching the caudal margin of the wings; the row 
of spines along the caudal margin of the abdominal segments not 
larger than the spines of the segment; ninth abdominal segment 
never with scattered spines larger than those of the other segments, 
b. Dorsum of the abdominal segments rugose and densely spinose on 

the first nine segments Euthisanotia Hubner. 

bb. Dorsum of abdominal segments rugose and moderately spinose 
on the first seven segments; the eighth and ninth segments 

roughened, but with very few spines Alypia Hubner. 

aa. Antennae never reaching the caudal margin of the wings; the row 
of spines along the caudal margin of the abdominal segments 
larger than those of the segment; ninth abdominal segment with 
scattered spines larger than those of the other segments. 

PsychomorpJta Harris. 
The following species were examined : 
Butkisanotia grata Fabricius, unto Hubner 
Alypia octomaculata Fabricius 
Psychomorpha epimems Drury 

Subfamily Acronyctinae 

The subfamily Acronyctinae as typified by the genus Acronycta 
has little to distinguish it from other subfamilies except that the cre- 
master is short and usually mound-like and the setae are always of 
the same size and length. With this genus there are here included 
several others, which probably do not form a natural group, though 
all possess cremastral setae of the same size and length and in all the 



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114 

members of the group only the prothoracic legs extend cephalaxi to 
the eye-pieces. All of these genera except Acronycta have prominent 
projections on the head and prothorax. The epicranial suture is 
present only in Eulonche. The labial palpi are well developed in all 
and the prothoracic femora are visible only in Eulonche, Acronycta, 
and Achatodes. The maxillae do not reach the caudal margin of the 
wings in any of the genera. The metathoracic legs are always visible, 
and in Eulonche the mesothoracic legs are adjacent on the meson. In 
Plusiodonta they extend to the caudal margin of the wings, but in 
none of the other genera. The antennae are usually equal in length 
to the mesothoracic legs and never reach the caudal margin of the 
wings. The genus Eulonche and a few species of Acronycta are very 
peculiar in that there are setae on the body arranged as in Arctiidae 
(Fig. 103). These are easily observed on the mesothorax and the 
parts of the abdomen where the sculpturing is not so dense. There 
are also a few short spines present on the tenth segment at the base 
of the cremaster. The genera of Acronyctinae may be separated 
as follows: 

a. Cephalic end of body with two large, rounded, rugose projections 
on the head, one on each side of the meson. 

b. Dorsum of abdomen very rugose on segments 1-7 except for a 
smooth caudal margin; groups of long setae present on thorax 

and abdomen; epicranial suture present Eulonche Grote. 

bb. Dorsum of abdomen never rugose, but with large lunate punc- 
tures on the cephalic third of segments 3-7; groups of long 
setae never present; epicranial suture absent. 

Achatodes Guen6e. 

aa. Cephalic end of body with a single median projection, or without 

projections. 

b. Cephalic end of body with a single median projection, either on 

the head or prothorax. 

c. A prominent projection on the body near the cephalic margin 

of the prothorax ; abdominal segments 1-3 with a broad smooth 

transverse ridge near the caudal margin; movable abdominal 

segments with thin flanged plates; cremaster very short and 

broad, with a short, stout, slightly curved seta inserted in each 

caudo-lateral angle Homopyralis Grote. 

cc. Projection always present on the head ; setae at the caudal end 
of the body always hooked, 
d. Distinct cremaster not present; a large sharp point at each 
caudo-lateral angle of the body, with some smaller points, 
and three short hooked setae inserted on each side : body dis- 
tinctly punctate, with a smooth band at the caudal margin 



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115 

of each segment; projection on the head blunt at the end 

and directed ventrad Phisiodonta GuenSe. 

dd. Distinct cremaster present, its caudo-lateral angles produced 

and pointed, with two hooked setae inserted on the ventral 

side of each ; body not punctate, slightly rugose ; projection 

on the head narrowed to a rounded point. .Anomis Hubner. 

bb. Cephalic end of body smooth, cremastral hooks slightly curved. 

Acronycta Treitschke. 
The following species were examined : 
Acronycta americana Harris, populi Riley, clarescens Guenee, hama- 

melis Guenee 
Eulonche oblinita Smith and Abbot 
Achatodes zeae Harris 
Homopyralis discalis Grote 
Plusiodonta compressipalpis Guenee 
Anomis erosa Hubner 

Subfamily Phytometrinae 

The members of this subfamily differ markedly from those of 
the other subfamilies of Noctuidae. The labrum is never in its nor- 
mal position but is located near the cephalic end of the body, while 
the wings and maxillae extend beyond the caudal margin of the fourth 
abdominal segment. The wings are produced into a sharp point near 
the meson of the ventral surface. The labial palpi are always visi- 
ble and a large portion of the prothoracic femur is exposed. The 
dorsal cephalic margin of the movable abdominal segments has a 
number of prominent furrows with slightly serrate ridges between. 
The prothorax is not as long as in the previously mentioned sub- 
families, being only two fifths the length of the mesothorax, and the 
caudo-lateral angles are somewhat produced. The dorsal surface of 
the body shows prominent grooves along the caudal margin of the 
metathorax and the first four abdominal segments. The edges of 
these grooves are somewhat serrate. The cremaster is somewhat 
cylindrical and rugose, with two long hooked setae and four shorter 
ones. The two genera studied are very closely related and may be 
separated as follows: 

a. Antenna reaching the caudo-lateral angle of the mesothoracic leg; 
cremaster with length and breadth approximately equal and longer 

than its longest setae Pkyiometra Haworth. 

aa. Antenna much shorter than the mesothoracic leg and never reach- 
ing its caudo-lateral angle; cremaster always longer than broad 
and about equal in length to its longest setae. 

Syngrctpha Hubner. 



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116 

The following species were examined: 
Phytometra brassicae Riley 
Syngrapha falcigera Kirby 

Subfamily Mominae 

The only genus studied in this group was Charadra. This re- 
sembles the Arctiidae very much more than it does most Noctuidae 
in the shape of the body, in the presence of setae arranged around 
the scars of larval verrucae, and in the absence of epicranial suture 
and visible labial palpi and femora of the prothoracic legs. The an- 
tennae are broader at the proximal end than is t)rpical in Noctuidae. 
The appendages are arranged more as they are in Noctuidae and 
there is a cremaster present, as long as the ninth and tenth abdom- 
inal segments, which bears hooked setae. The only known Arctiidae 
which have long cremasters are provided with flanged plates on the 
movable abdominal segments and the cremastral setae are never 
hooked. The prothoracic leg extends cephalad between the sculptured 
eye-piece and the antenna, but the mesothoracic leg never reaches 
the eye-pieces. The body is slightly punctate along the cephalic mar- 
gin of the movable abdominal segments. The pupae are found in 
thin silken cocoons, which differ from those of the species of 
Arctiidae in that none of the larval hairs are used in their construc- 
tion. 

The following species was the only one examined : 
Charadra deridens Guenee. 

Subfamily Hypeninae 

The only genus available for study of this group as given in 
Dyar's list was Plathypena, but as Balsa possesses the pupal charac- 
ters which distinguish this from other subfamilies, it is included in 
the Hypeninae for the present. These characters are the presence 
of two long and six short hooked setae at the caudal end of the body 
and the fact that the prothoracic legs and the long point of the meso- 
thoracic legs extend cephalad to the eye-piece, or, as in Balsa, to the 
maxillary palpus, which is always present in this genus. Plathypena 
shows the epicranial suture, but it is not found in Balsa. Both genera 
have the spiracles slightly produced, and in Plathypena they are on 
small elevations. The labial palpi are present in both, but the femora 
of the prothoracic legs are visible only in Balsa. The genera may be 
separated as follows: 



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a. Dorsum of prothorax, metathorax, and first three abdominal seg- 
ments with a distinct median ridge; antennae and mesothoracie 
legs reaching the caudal margin of the wings and longer than the 
maxillae ; caudal margin of mesothorax never with a row of shal- 
low depressions Plcpthypena Grote. 

.aa. Dorsum of body never with a median ridge; antennae and meso- 
thoracie legs not reaching the caudal margin of the wings and not 
longer than the maxillae ; caudal margin of mesothorax with a row 
of shallow depressions Baha Walker. 

The following species were examined : 
Plathypena scabra Fabricius 
Balsa malana Fitch 

Subfamily Catocalinae 

This group is distinguished from all other noctuids by the pres- 
<^nce of a whitish "bloom" on the surface of the pupa, which is re- 
tained even in alcoholic specimens. Both prothoracic and meso- 
thoracie legs extend cephalad to the eye-pieces. The labial palpi are 
always present, but the femora of the prothoracic legs are seldom 
visible. The epicranial suture is found throughout the genera Catoc- 
ala and Eunetis but is lacking in the remainder of the subfamily. 
The antennae, mesothoracie legs, and maxillae either reach the cau- 
<lal margin of the wings or very closely approach it. The cremaster 
is usually very short or absent, and the setae at the caudal end of the 
body are usually of two sizes and inserted at different levels except 
in the genus Eunetis. This generic name is applied to certain species 
of the genus Catocala of Dyar's list which have a short cremaster, 
slightly broader at the caudal end, bearing about eight slightly 
oirved setae which are usually directed towards the meson. The fol- 
lowing table will serve to separate the genera of Catocalinae: 

A. Epicranial suture present; body tapering rapidly from the fifth ab- 
dominal segment so that it is more slender in appearance than the 
typical noctuid, the lateral margins of abdominal segments 8-10 as 
seen in dorsal view convergent and not strongly convex, 
b. Cremaster, if present, very short and narrowed at the caudal end, 
and with eight long hooked setae of two sizes, some larger and 

more heavily chitinized than the others Catocala Schrank. 

bb. Cremaster broadened at the caudal end, usually with eight short, 
stout setae which are slightly curved and usually directed tow- 
ards the meson Eunetis Hiibner. 

^Wi. Epicranial suture not present; body of typical noctuid shape, with 
the lateral margins of abdominal segments 8-10 distinctly convex 
as seen in dorsal view. 



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b. Head with a distinct tubercle near the base of each antenna. 

EupartTienos Grote. 
bb. Head without a distinct tubercle at the base of each antenna, 
c. Thorax and appendages with deep indeterminate transverse 
striations; median caudal spines of cremaster somewhat en- 
larged near the tip ; metathoracic legs never present. 

Pheocyma Hiibner. 

cc. Thorax and appendages approximately smooth ; median caudal 

spines of cremaster never enlarged near the tip ; metathoracic 

legs always apparent caudad of the maxillae. 

d. Cremaster with spines practically all of the same size, no two 

being larger and longer than the others. .Caenurgia Walker. 

dd. Cremaster with two spines larger and longer than the others. 

Zale Hiibner. 
The following species were examined : 
Euparthenos nuhilis Hubner 
Pheocyma lunifera Hubner 

Caenurgia erechtea Cramer, crassiuscula Haworth 
Zale ccUycanthata Smith and Abbot, lunata Drury 
Catocala illecta Walker, unijuga Walker, briseis Edwards, verecunda 
Hulst, aholibah Strecker, ilia Cramer, innubens Guenee, neo- 
gania, Smith and Abbot, pacta Linnaeus, sponsa Linnaeus 
Bunetis blandula Hulst, ultronia Hiibner, grynea Cramer 

Subfamily Sarrothripinae 

This group is readily distinguished because it has neither ore- 
master nor setae at the caudal end of the body, which is probably 
due to the fact that its members are found in thick cocoons. The 
dorsal surface of the body is very irregularly rugose with spinous 
elevations, and there is a distinct row of spines along the caudal mar- 
gin of the fifth abdominal segment extending from the rugose area on 
the dorsum around nearly to the meson of the ventral surface. A 
few spines are present in a similar position on the fourth abdominal 
segment. The epicranial suture is always present, the labial palpi are 
visible for their entire length, and only a small portion of the pro- 
thoracic femur is exposed, or it may be entirely concealed. The 
maxillae never reach the caudal margin of the wings, being about 
seven eighths of their length, with the mesothoracic legs meeting 
just caudad of them. The antennae always reach to the caudal mar- 
gin of the wings, while the mesothoracic legs do not, but the latter 
are slightly longer than the maxillae. Both prothoracic and meso- 
thoracic legs extend cephalad to the eye-piece, the mesothoracic legs 
extending between the sculptured eye-piece and the antenna. 



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The members of this subfamily have been treated as the family 
Nycteolidae by some authors, but there is no evidence in the pupae 
to separate them from the family Noctuidae. 

The following species were examined : 
Sarrothripus revayana Scopoli, proteella Dyar. 

Family Arciudae 

The members of this family all possess distinct setae arranged 
around the scars of the larval verrucae. These setae are seldom con- 
spicuous enough to be seen with the naked eye as in Liparidae, but 
are easily visible with the aid of the microscope. The labial palpi are 
never visible, unless as small triangular areas caudad of the labrum, 
except in Halisidota, where they are exposed for their entire length. 
The femora of the prothoracic legs are never visible. The shape of 
the body is characteristic, being slightly concave on the dorsum in 
the region of the metathorax (Fig. 104). Certain genera of Noctuidae, 
Acronycta, Eulonche, and Charadra, also show setae arranged around 
the scars of the larval verrucae. In the two genera first named, both 
labial palpi and prothoracic femora are exposed, while Charadra pos- 
sesses a long cremasttr bearing hooked setae. Those genera of 
Arctiidae with a cremaster never have hooked setae, but all cremastral 
setae are flattened at the distal end. The epicranial suture is never 
present in any member of this family. The prothorax is usually long, 
often half the length of the mesothorax as in most Noctuidae. The 
genus Ctenucha, included with the S)mtomidae in Dyar's list, shows no 
dharacters to distinguish it from the Arctiidae and it is probable that 
other genera of this family should be included here. The genera of 
Arctiidae may be separated as follows : 

a. Abdominal segments 5-7 never with a flanged plate along the ce- 
phalic margin, or with deep furrows between these segments when 
the body is retracted ; maxillae nearly as long as the wings ; mesotho- 
racic wings never meeting on the meson caudad of the appendages, 
b. Dorsal surface of abdomen flattened; body broadly rounded at 
caudal end and bearing a row of short setae which are slightly 
curved at tip ; body brown, concolorous. 
c. Labial palpi showing for about one sixth of the length of the max- 
illae; body 18-20 mm. in length; dorsal surface of abdomen 

never with depressed areas Halisidota Hiibner. 

cc. Labial palpi never showing except as a small triangular piece 
about a millimeter in length ; body 12-15 mm. in length ; dor- 
sal surface of abdomen with depressed areas on meson of each 
segment at cephalic margin and one on each side adjacent to the 
spiracles Euchaetias LymaM. 



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bb. Dorsal surface of abdomen not flattened ; body tapering at caudal 
end ; color yellowish or chestnut-brown, strikingly marked with 
black. 

c. Distinct cremaster present Haploa Hiibner. 

cc. Distinct cremaster never present, 
d. Antennae about seven eighths the length of the wings ; maxiUae 
never reaching the caudal margin of the wings. 

Uietkeisa Hiibner. 
dd. Antennae as long as the wings ; maxillae always reaching the 

caudal margin of the wings Cienucha Kirby. 

aa. Abdominal segments 5-7 with a flanged plate along the cephalic 
margin, and with deep furrows between the movable segments 
when the body is retracted ; maxillae never as long as the wings, 
usually about two thirds the length; mesothoracic wings always 
meeting on the meson caudad of the appendages, 
b. Abdominal segments 4-6 with a similar flanged plate adjacent to 
. the caudal margin of the segment, 
c. Appendages, and usually the thorax, roughened with indetermi- 
nate transverse striations ; abdominal segments densely, coarsely 
punctate, 
d. Head with a small tubercle at the proximal end of each an- 
tenna; antennae always much shorter than the prothoracic 

legs Estigmene Hiibner. 

dd. Head without a tubercle at the proximal end of each antenna ; 
antennae as long as, or longer than, the prothoracic legs, 
e. Body usually 18-20 mm. in length, stout; setae of the cre- 
master of various sizes and lengths, the shortest ones only 
about half the length of the longest, and irregularly ar- 
ranged, always fifteen or more in number. 

Diacrisia Hiibner. 

ee.. Body seldom over 12 mm. in length, varying from 10-15 

mm., rather slender ; setae of the cremaster of practically 

the same size and length, arranged in two transverse rows, 

with four in the dorsal and eight in the ventral row, never 

as many as fifteen in number Hyphantria Harris. 

cc. Thorax and appendages smooth and polished; body sparsely, 

finely punctate Ecpantheria Hubner. 

bb. Abdominal segments 4-6 never with a flanged plate adjacent to 
the caudal margin, 
c. Distinct cremaster always present and long, bearing setae at the 
caudal end; antennae about equaling the maxillae in length, 
usually three fourths the length of the wings. 

Apantesis Walker, 
cc. Distinct cremaster never present ; a row of setae at the caudal 
end of the body ; antennae very much shorter than the max- 
illae, being about half the length of the wings Isia Walker. 



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121 

The following species were examined: 
Halisidota iessellaris Smith and Abbot, caryae Harris 
Euchaetias egle Drury 
Haploa clymene Brown 
Utetheisa bella Linnaeus 
Ctenucha virgitiica Charpentier 
Bstigmene acraea Drury 
Diacrisia virginica Fabricius 
Hyphantria cunea Drury 
Bcpantkeria deflorata Fabricius 
Apantesis virgo Linnaeus, michabo Grote, arge Drury, phyllira 

Drury, nais Drury 
Isia isabella Smith and Abbot 

Family LiPARroAE 

This family, like the Arctiidae, is characterized by the presence 
of setae arranged around the scars of the lan^al verrucae. In the 
Liparidae the setae are long and coarse, and easily visible to the un- 
aided eye. With the exception of Porthetria all the genera examined 
show a characteristic arrangement of appendages (Fig. 105). In Por- 
thetria the labial palpi were usually concealed by the maxillae, al- 
though a large number of pupae show them present, as in Figure 
105. The epicranial suture is never present. The maxillae are always 
short, never more than two fifths the length of the wings. The legs 
are usually shorter than in most pupae, the mesothoracic legs never 
reaching the caudal margin of the wings. The antennae are pectinate 
and are longer and broader in the male than in the female. The cre- 
master is always present, smooth, longer than broad, and bears short 
hooked setae at the distal end. 

Most of the species examined show a remarkable uniformity of 
characters, and considerable difficulty was encountered in separating 
the genera. The difficulty lies in the fact that there is considerable 
difference between the sexes, not only in the length and breadth of 
the antennae, but in the size and arrangement of other appendages. 
In Hemerocampa the adult females are apterous, and the wings in 
the pupa are not as long as in the males. The wings of the females 
reach slightly over the cephalic margin of the fourth abdominal seg- 
ment, while in the male they reach to the caudal margin of that seg- 
ment. In Hemerocampa the dorsum of the first three abdominal seg- 
ments is covered on each side of the meson with small vesicles. The 
following table will serve to separate the genera of Liparidae : 



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a. Spiracular furrows never present on the cephalic margin of the mov- 
able abdominal segments; labial palpi present and well developed; 
long setae not present on the face-parts near the caudal margin of 
the head, on the clypeus, or on the sculptured eye-pieces, 
b. Labial palpi about equal in width to each maxilla; cremastral 
setae about one third the length of the cremaster. 
c. Dorsal surface of abdomen densely covered with long setae ; body 
brown, concolorous except sometimes for the lighter transverse 
conjunctiva ; antennae with the stem of the flagellum very broad 
and distinctly elevated, 
d. Abdominal segments 8-10, in dorsal view, distinctly narrowed 
and tapering; clypeus and labrum on a level with the other 
face-parts; cremaster directed caudad; antennae of male 
reaching about half the distance along the mesothoracic legs, 

and distinctly pointed DasycTiira Stephens. 

dd. Abdominal segments 8-10, in dorsal view, not narrowed and 
distinctly tapering; clypeus and labrum both elevated above 
the level of the other face-parts ; cremaster directed dorsad ; 
distal half of the antennae of the female distinctly narrowed 
and pointed, reaching about half the distance along the 

mesothoracic legs Olene HQbner. 

cc. Dorsal surface of abdomen not very densely covered with setae ; 
body white variously marked with brown, or vice versa ; anten- 
nae in both sexes with the stem of the flagellum scarcely indi- 
cated and the distal end always rounded, in the male extend- 
ing about half the length of the wings and curved mesad so that 
they often meet, in the f eihale never reaching half the distance 
along the mesothoracic legs and never meeting on the meson. 

Hemerocampa Dyar. 
bb. Labial palpi only half the width of each maxilla; cremastral 
setae about half the length of the cremaster; antennae in the 
male broad for almost their entire length and extending about 
three fourths the length of the wings, adjacent or meeting on the 
meson, in the female pointed at the distal end and reaching 
about half the distance along the mesothoracic legs. 

Euproctis Hiibner. 
aa. Spiracular furrows always present on the cephalic margin of the 
movable abdominal segments and extending almost to the meson, 
five or six in number and separated by sharply carinate ridges; 
caudal portion of face-parts, clypeus, and sculptured eye-pieces 
with coarse setae similar to those on the body, labial palpi often con- 
cealed, but sometimes visible as in the remainder of the family. 

PortTietria Hiibner. 
The following species were examined: 
Dasychira pudibunda Linnaeus 
Olene manto Strecker 



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Hemerocampa leucostigma Smith and Abbot 
Porthetria dispar Linnaeus 
EuprocHs chrysorrhoea Linnaeus 

SuPERPAMiLY BOMBYCOIDEA 

This superfamily includes those families in which the body is 
more or less densely covered with setae and which usually retain 
the labial palpi. They seem to be more nearly related to the Satur- 
nioidea than to any other superfamily, although the Lasiocampidae 
show certain points of relationship with the Liparidae. All the 
members of the superfamily, so far as known, are found in thick 
silken cocoons, much like those of the Saturniidae. The families of 
Bombycoidea may be separated as follows : 

a. Epicranial suture present; labial palpi visible Lasiocampidae. 

aa. Epicranial suture never present ; labial palpi concealed by the maxil- 
lae except for a small triangular area at the proximid end. 

BOMBYCIDAE. 

Family Lasiocampidae 

The members of this family usually have mouth parts and appen- 
dages arranged as in the Liparidae (Fig. io6). The epicranial su- 
ture is always present and the vertex is longer than that found in any 
but the more generalized forms. The maxillae are short, never more 
than one third the length of the wings, and extend very slightly be- 
yond the distal ends of the labial palpi, or may be shorter than the 
palpi. 

The antennae are broad and pectinate, and never extend as far 
caudad as the prothoracic legs. The coxae of the prothoracic legs and 
sometimes of the mesothoracic pair are often visible caudad of the 
maxillae and labial palpi. The prothoracic legs are slightly more than 
lialf the length of the wings and the mesothoracic legs never reach 
the caudal margin of the wings. The setae of the body are very con- 
spicuous except in Tolype, but are not arranged around the scars of 
the larval verrucae. The movable segments are capable of being re- 
tracted till only their caudal margins are visible. There is never a 
cremaster present, and there are no hooked setae at the caudal end 
of the body. The body is broadly rounded at the caudal end and 
the body setae are usually a little longer and coarser in this region. 
This family is considered by many authors to be more specialized 
than any of the Saturnioidea, but the presence of the epicranial su- 
ture and exposed labial palpi shows that they are much more gener- 
alized. The genera of Lasiocampidae may be separated as follows : 



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124 

a. Entire surface of body except the appendages with a dense covering- 
of fine short setae, giving it a furred appearance; abdominal seg- 
ments 8-10 considerably narrower than the remainder of the ab- 
domen Malacosoma Hiibner. 

aa. Entire surface of body except appendages never with a dense cover- 
ing of setae giving it a furred appearance, either with a very 
sparse covering, or without visible setae; abdominal segments 8-^ 
10 not noticeably narrower than the remainder of the abdomen, 
b. Body sparsely covered with very fine short setae except at the cau- 
dal end, where they are longer and coarser ; tenth segment broadly 
rounded at the caudal end. 
c. Lateral surface of abdomen on either side of the spiracles with 
setae much thicker than on the dorsum ; dorsal surface of abdo- 
men very coarsely punctate Lasiocampa Schrank. 

cc. Lateral surface of abdomen on either side of the spiracles never 
with the setae more numerous than on the dorsum ; dorsal sur- 
face of abdomen very finely punctate Cosmotricke Hiibner. 

bb. Body without any visible covering of setae; tenth segment 
abruptly narrowed at the caudal end, suggesting a cremaster. 

Tolype Hiibner, 
The following species were examined: 
Malacosoma americana Fabricius, disstria Hiibner 
Lasiocampa querelas Linnaeus (Europe) 
Cosmotricke potatoria Linnaeus (Europe) 
Tolype velleda StoU 

Family BoMBYcroAE 

This family includes the genus Bombyx, which is domesticated in 
various parts of the world (Fig. 107). The body is covered with 
rather coarse short setae which are somewhat longer at the caudal 
end of the body. The epicranial suture is never present. The ap- 
pendages are arranged much as in Lasiocampidae. The labial palpi 
are almost concealed, being overlaid by large ovate appendages, ap- 
pearing to be heavily veined, which are presumably the maxillae (no 
dissections were made of this species). The mandibles are repre- 
sented by strongly elevated tubercles. The coxae of one pair of legs, 
probably the mesothoracic, are adjacent on the meson caudad of the 
maxillae. The legs are short, and both prothoracic and mesothoracic 
pairs lie adjacent on the meson. The antennae are pectinate, broad 
at the proximal end and rapidly narrowed, ending in a point oppo- 
site the distal ends of the prothoracic legs. The mesothoracic wings 
lie adjacent on the meson caudad of the mesothoracic legs. The 
movable segments may be retracted so that only their caudal margins 
are visible. There is no cremaster present. The pupa strongly re- 



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125 

sembles those of certain species of Saturniidae and it is quite probable 
that they had a common ancestor, although the Bombycidae are un- 
doubtedly more generalized. The only species in America is Bom- 
byx mori Linnaeus. 

SuPERPAMiLY NOTODONTOIDEA 

The families included here never have the entire labial palpi ex- 
posed, but a very small triangular or polygonal area is sometimes visi- 
ble just caudad of the labrum. Some genera of Geometridae have the 
prothoracic femora exposed, while the epicranial suture is present in 
some of the members of each family. Although the larvae of Geomet- 
ridae are easily recognized and are very readily distinguished from 
those of the Notodontidae, the pupae show much closer relationships, 
and it is difficult to draw a hard and fast line between the two fami- 
lies. The three families included here have probably had a common 
ancestor, and although the Dioptidae retain the epicranial suture they 
must be considered the most specialized, as both Geometridae and 
Notodontidae show more generalized characters in some of their gen- 
era. The following table may be used to separate the families of 
Notodontoidea: 

a. Antennae never extending beyond the caudal margin of the wings ; 
dorsum of abdomen never with a prominent hooked seta on each side 
of the meson of segments 7-10. 

b. Maxillae usually more than three fifths the length of the wings, if 
not, then the caudal end of the body with hooked setae, or 
the spiracles of the third abdominal segment concealed by the 
wings and those of the sixth segment farther ventrad than those 
of the other segments ; prothoracie femora often exposed ; a deep 
furrow usually present on the dorsum of the abdomen between 
the ninth and tenth segments ; caudal margin of mesothorax never 
with a row of deep pits with smooth tubercle-like areas between. 

Geometrtoae. 
bb. Maxillae seldom exceeding three fifths the length of the wings, if 
so, then the caudal margin of the mesothorax with a row of deep 
pits with smooth, elevated, quadrangular tubercle-like areas be- 
tween them ; or with the entire body surface coarsely punctate ; 
abdominal spiracles of the third segment never concealed by the 
wings, and those of the sixth never farther ventrad than the re- 
mainder; prothoracic femora never exposed; a furrow never 
present on the dorsum of the abdomen between segments 8 and 9 
except in Datana, where the cremaster is of the type shown in 
Figure 112 Notodonttoae. 



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aa. Antennae extending beyond the caudal margin of the wings ; dorsum 
of abdomen with a prominent hooked seta on each side of the meson 
of segments 7-10 DioprmAE. 

Family Geometrtoae 

The pupae in this family (Figs. io8 and 109) never have the labial 
palpi exposed except as small triangular or polygonal areas caudad 
of the labrum. The maxillary palpi are, never present. The epicranial 
suture is present in some of the genera in Groups A and C. Some of 
the genera have the femora of the prothoracic legs exposed. In all 
the genera examined, either the prothoracic or mesothoracic leg, or 
sometimes both, extended cephalad between the sculptured eye-piece 
and the antenna. The prothoracic and mesothoracic legs are longer 
than is usual in most families, the former being usually three fourths 
the length of the wings, while the latter always extend to the caudal 
margin of the wings. Many of the genera show the f ronto-cl)rpeal su- 
ture extending from the proximal ends of the antennae and directed 
caudad towards the invaginations for the anterior arms of the ten- 
torium. The suture is very distinct for the cephalic part and is often 
indicated by a slight furrow for the remainder of the distance. In 
the genus Haematopsis there is a prominent cephalic projection 
bearing hooks which hold the suspensory threads, as this pupa is not 
found in a cocoon. The antennae vary little throughout the family, 
and are generally equal in width to, or wider than, the prothoracic 
legs, usually extending to the caudal margin of the wings. The meso- 
thoracic wings usually extend farther caudad than in the nearly related 
families, reaching almost to the caudal margin of the fourth abdom- 
inal segment, although not visible in ventral view. The mesothorax is 
very short in some genera, particularly in those of Group D, where it 
is never twice the length of the prothorax. The mesothoracic spir- 
acles very often have a decided projection adjacent to their caudal 
margin, which is usually flattened or tuberculate in form and often 
covered with very fine short setae. The abdominal spiracles are some- 
times produced, and in nearly all genera the sixth spiracle, and some- 
times the seventh also, is considerably farther ventrad than the re- 
mainder. The abdomen is usually coarsely punctate, sometimes rough- 
ened with deep transverse striations. A cremaster of some kind is al- 
ways present. One of the most interesting structures of the abdomen is 
the dorsal furrow between the ninth and tenth segments. This is 
usually deep and fringed with very fine setae. This furrow fre- 
quently projects caudad on the lateral surface of the body and this 
portion is often separated from the dorsal furrow. This dorsal fur- 



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127 

row is present in all members of Group A except Ennomos and in a 
few members of Group D. These dorsal furrows are found in other 
families, notably in the Gelechiidae and some Pyralididae. 

The attempt to classify the pupae of Geometridae was seriously 
hampered by lack of material. Reared material was very hard to ob- 
tain, as the larvae develop slowly and seem to be very susceptible to 
disease. The available material, moreover, did not seem to fall in 
with any of the existing schemes of classification, so that the only 
practical solution of the difficulty was to divide them into groups ac- 
cording to the pupal characters. These may or may not be natural 
groups, but they will serve to indicate relationships in some degree. 
According to the pupal characters there seem to be two large divi- 
sions, one with hooked setae on the cremaster, the other without them. 
As the presence of hooked setae is a more generalized character, 
the groups in which they are present should be the more generalized, 
and the presence of the epicranial suture strengthens this view. 
Group A includes representatives of the subfamilies Sterrhinae, En- 
nominae, and Hydriomeninae as listed by Dyar. Group B includes 
for the present only the genus Haematopsis. Group C includes the 
genera Alsophila and Brephos, which are similar in many respects and 
possess the same type of cremaster. This must also be considered as 
a generalized group since the epicranial suture is present in Alsophila. 
.Brephos has usually been considered as the most generalized geomet- 
rid, but no epicranial suture has been located. The maxillae are 
also much shorter in Alsophila. In Group D the epicranial suture is 
never present. Spiracular furrows are frequently found in Groups 
A and D. The adult females of some of the geometrid species are 
apterous. Although abundant material of one such species, Paleac- 
rita vernata, was examined, the pupal wings were found as well 
developed in the female as in the male. These groups of Geometridae 
may be separated as follows: 

a. Cremaster with hooked setae. 

b. Setae of the cremaster always of two sizes, usually either the two 
or four farthest caudad much larger than the others ; dorsum of 
abdomen usually with a deep furrow between abdominal seg- 
ments 9 and 10; body never with a bifurcate projection at the 

cephalic end Group A. 

bb. Setae of the cremaster always of the same size ; dorsum of abdo- 
men never with a furrow between segments 9 and 10 ; body with 
a long bifurcate projection at the cephalic end, densely covered 
with hooked setae Group B. 



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128 

aa. Cremaster never with hooked setae. 

b. Cremaster a stout T-shaped spine Group C. 

bb. Cremaster always bifurcate Group D. 

Group A 

This group includes species in which the cremastral setae are of 
two sizes, and which generally show the epicranial suture, and a dor- 
sal furrow between the ninth and tenth abdominal segments (Fig. 
109). Spiracular furrows are also frequently present. The genera 
included in this group may be separated as follows : 

a. Epicranial suture always present; prothoracic femora always ex- 
posed, usually a large portion; spiracular furrows never present; 
dorsum of abdomen with a distinct furrow between segments 9 and 
10, and also on the lateral surface of segment 10. 
b. Dorsum of fifth abdominal segment with a distinct furrow on the 
cephalic margin ; caudal margin of the furrow between the ninth 
and tenth abdominal segments with finely serrate edges and with- 
out setae Hydria Hiibner. 

bb. Dorsum of fifth abdominal segment never with a distinct furrow 

on the cephalic margin; caudal margin of the furrow between 

the ninth and tenth abdominal segments coarsely serrate. 

c. Prothoracic and mesothoraeic legs extending cephalad between 

the sculptured eye-piece and the antenna, the mesothoraeic leg 

extending almost as far cephalad as the prothoracic; caudd 

margin of the furrow between the ninth and tenth abdominal 

segments without setae ; cremaster distinctly constricted at the 

proximal end and circiilar in outline Eois Hiibner. 

cc. Prothoracic leg extending cephalad between the sculptured eye- 
pieces and the antenna, the mesothoraeic leg never reachhig 
farther cephalad than the caudal margin of the eye-piece ; cau- 
dal margin of the furrow between the ninth and tenth ab- 
dominal segments usually fringed with fine setae; cremaster 
not constricted at the proximal end, triangular in outline, 
d. Cremaster with hooked setae of nearly the same size, the me- 
dian caudal setae being only slightly larger than the others ; 
three setae inserted along each side of the cremaster, the other 
two cephalad of these and slightly mesad. 

Tephroclystis Hiibner. 
dd. Cremaster with the hooked setae of widely differing sizes, the 
two median caudal setae being much larger and longer than 
the others, a smaller seta adjacent on each side, while the 
other setae are arranged in a tranverse row just cephalad 
of the caudal setae Cinglis Guen6e. 



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aa. Epicranial suture never present. 

b. Prothoracie femora exposed, but only a very narrow portion, 
e. Spiracular furrows present on the fifth abdominal segment ; ven- 
tral surface of head with a prominent transverse ridge extending 
from, about the middle of the glazed eye-piece ; body setae arising 
from small dark brown or black papillae. .PhUobia Duponchel. 
cc. Spiracular furrows never present on the fifth abdominal seg- 
ment ; ventral surface of head never with a transverse ridge ; 

body setae arising from small pits Sabulodes Guen^e. 

bb. Prothoracie femora never exposed, 
c. Antennae distinctly elevated and covered with five or six rows of 
small round tubercles ; distinct ridges or flanged plates present 
along both margins of the movable segments, 
d. Cremaster with the two median caudal spines very much larger 
than the others; distinct furrow always present between the 
ninth and tenth abdominal segments; color dark brown. 

Nacophora Hulst. 

dd. Cremaster with the four caudal spines about the same size and 

much larger than the others ; distinct furrow never present 

between the ninth and tenth abdominal segments; color 

usually white, sometimes partly brown. 

Ennomos Treitschke. 

cc. Antennae elevated but never with distinct rows of tubercles ; 

ridges or flanged plates never present on the movable segments. 

d. Maxillae about two thirds the length of the wings ; prothoracie 

mesothoracic, and metathoracic legs all meeting on the meson 

caudad of the maxillae Xanthotype Warren. 

dd. Maxillae always more than two thirds the length of the wings ; 
prothoracie and mesothoracic legs never meeting on the 
meson caudad of the maxillae, 
e. Spiracular furrows present on the cephalic margin of the 
fifth abdominal segment, with more or less interrupted 
ridges between ; cremaster with the four caudal setae always 

larger than the others Ania Stephens. 

ee. Spiracular furrows never present on the cephalic margin 
of the fifth abdominal segment, cremaster with two caudal 
setae larger than the others CingUia Walker. 

Group B 

The pupae of this group are easily distinguished by the long bifur- 
cate projection at the cephalic end of the body, which is covered with 
hooked setae. There are never spiracular furrows present on the ce- 
phalic margin of the movable segments. The mesothoracic legs meet on 
the meson caudad of the maxillae. The body is very slender and never 
punctate. This group includes the genus Haematopsis. 



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130 

Group C 

The species of this group are distinguished by the peculiar T- 
shaped cremaster. The genera may be separated as follows : 

a. Epicranial suture present; prothoracic and mesothoracic legs meeting 

on the meson caudad of the maxillae AlsophUa Hiibner. 

aa. Epicranial suture never present; prothoracic and mesothoracic legs 
never meeting on the meson caudad of the maxillae. 

BrepJios Ochsenheimer. 

Group D 

This group is characterized by the presence of a bifurcate cre- 
master. The epicranial suture is never present, but a portion of the 
prothoracic femora is exposed in many genera. The following table 
will serve to separate the genera of this group : 

a. Prothoracic femora visible. 

b. Cephalic margin of the fifth abdominal segment with one deep 

spiracular furrow with strongly chitinized edges, just cephalad 

of each spiracle. 

c. Mesothoracic spiracle with a broad, very strongly elevated ridge or 

oval tubercle adjacent to its caudal margin which is covered with 

fine short setae; surface of spiracular furrow with distinct 

punctures Physostegania Warj:en. 

cc. Mesothoracic spiracle with only a very narrow, slightly elevated 
ridge adjacent to its caudal margin, usually covered with setae ; 
surface of spiracular furrow without distinct punctures, 
d. Dorsal surface of abdomen never with a distinct furrow be- 
tween the ninth and tenth segments, its caudal margin finely 
serrate; surface of the spiracular furrow almost smooth. 

Ectropis Hiibner. 
d. Dorsal surface of abdomen never with a distinct furrow between 
the ninth and tenth segments ; surface of the spiracular fur- 
rows strongly rugose Cleora Curtis. 

bb. Fifth abdominal segment without any deep spiracular furrow; 
cephalic margin of the segment deeply punctate, the punctures 
sometimes confiuent; mesothoracic spiracle with a broad 
strongly elevated ridge or tubercle adjacent to its caudal mar- 
gin; dorsal surface of abdomen without a distinct furrow be- 
tween the ninth and tenth segments Cymatophora Hiibner. 

aa. Prothoracic femora never visible; deep spiracular furrows always 

present on the cephalic margin of the fifth abdominal segment; 

mesothoracic spiracle always with a prominent elevation adjacent 

to its caudal margin. 

b. Dorsal surface of abdomen never with a furrow between the ninth 

and tenth segments; abdominal spiracles very strongly produced; 



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131 

cremaster often showing two lateral setae on each side near the 
proximal end. 
bb. Dorsal surface of abdomen with a distinct furrow between the 
ninth and tenth segments, its caudal margin coarsely serrate ; a 
prominent lateral depression or furrow present on the lateral 
surface of the tenth abdominal segment, 
c. Maxillae four fifths the length of the wings or less ; mesothoracic 
legs meeting on the meson just caudad of the maxillae; an- 
tennae more than twice as wide at the proximal as at the distal 

end .Lycia Hiibner. 

cc. Maxillae always more than four fifths the length of the wings ; 
mesothoracic legs never meeting on the meson caudad of the 
maxillae; antennae of almost the same width throughout, 
never twice as wide at the proximal as at the distal end. 

Erannis Hiibner. 

The following species were examined : 
Group A 

Hydria undulata Linnaeus 

Eois inductata Guenee 

Tephroclystis interruptofasciaia Packard, absinthiaia Clerck 

Cinglis similaria Walker 

Philobia enotata Guenee 

Sabulodes lorata Grote, transversata Drury 

Nacophora quemaria Smith and Abbot 

Hnnomos subsignarius Hiibner, magnarius Guenee 

Xanthotype crocataria Fabricius 

Ania limbata Haworth 

Cingilia catenaria Drury 

Cosymbia serrulata Packard 
Group B 

Haematopsis grataria Fabricius 
Group C 

Alsophila pometaria Harris 

Brephos infans Moschler 
Group D 

Physostegama pusttUaria Guenee 

Bctropis crepuscularia Denis and Schiffermueller 

Cleora pampinaria Guenee 

Cymatophora ribearia Fitch 

Paleacrita vernata Peck 

Lycia cognataria Guenee 

Brannis tiliaria Harris 



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Family Notodontidae 

The pupae of this family never show more than a small triangular 
or polygonal proximal portion of the labial palpi, and maxillary palpi 
are never present. The femora of the prothoracic legs are never ex- 
posed. The epicranial suture is present in the genera Apatelodes and 
Melalopha. The maxillae never reach the caudal margin of the wings. 
The antennae are always widest at their proximal ends, and there the 
width exceeds the greatest width of the prothoracic legs. Each an- 
tenna tapers gradually to a pointed tip and the tips often lie adjacent 
on the meson caudad of the other appendages. The metathoracic legs 
are seldom visible. The mesothoracic leg never reaches cephalad to 
the eye-pieces. The abdomen is always punctate and in most species 
the punctures are large. A cremaster is usually present and there are 
various types, as in Figures iii, 112, 113. Packard divided the Noto- 
dontidae into six subfamilies. The pupae examined show that these 
subfamilies are well founded, but only tables to genera are given here 
as so few species of Notodontidae were examined. The genera 
Schizura and Heterocampa are not well defined and the species are 
separated with difficulty. The species are listed, however, under the 
subfamily name. 

Some authors believe that the genus Apatelodes belongs to the 
European f amjly Eupterotidae, and is incorrectly listed with the Noto- 
dontidae. As no pupae of Eupterotidae have been examined, it is im- 
possible to say whether pupal characters would justify this change. 
There are, however, no pupal characters as far as observed, which 
would prevent its being included with the Notodontidae. The two 
species differ widely and are possibly not congeneric. The following 
tables will serve to separate the genera of Notodontidae : 

a. Maxillae one third, or less, the length of the wings ; both prothoracic 
and mesothoracic legs meeting on the meson caudad of the maxillae ; 
abdomen very finely punctate. 

b. Thorax and abdomen thickly covered with very fine, short setae ; 
cremaster a stout spine about one millimeter in length with two 
short recurving hooks at the tip, each of which bears two or more 

very fine setae Melalopha Hiibner. 

bb. Thorax and abdomen never thickly covered with fine, short setae ; 
cephalic margin of first abdominal segment without tubercles; 
cremaster never as described above, sometimes absent, 
c. Abdominal segments 2-7 with distinct flanged plates at both 
cephalic and caudal margins, the cephalic plate interrupted by 
deep pits, giving it the appearance of a row of square tuber- 
cles ; appendages not at all elevated, making a smooth even 



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surface; cephalic end of body not elevated between the an- 
tennae ; short cremaster sometimes present. 

Apatelodes Packard, 
cc. Abdominal segments 2-7 never with flanged plates; appen- 
dages distinctly elevated ; cephalic end of body elevateid be- 
tween the antennae ; cremaster never present. 

Harpyia Ochsenheimer. 
aa. Maxillae always more than one third the length of the wings ; never 
with both prothoracic and mesothoracic legs meeting on the meson ; 
abdomen usually rather coarsely punctate, 
b. Maxillae from one half to three fifths the length of the wings; 
mesothoracic legs meeting on the meson caudad of the maxillae ; 
appendages roughened with deep indeterminate striations; ab- 
dominal segments coarsely punctate; a distinct, deep furrow on 
• the dorsum between the ninth and tenth abdominal segments; 

cremaster short, bifurcate, each half with several short spiny 

processes, usually directed laterad Datana Walker. 

bb. Maxillae more than three fifths the length of the wings ; neither 
prothoracic nor mesothoracic legs meeting on the meson caudad 
of the maxiUae; appendages usually with shallow indetermi- 
nate striations; a distinct furrow never present on the dorsum 
between the ninth and tenth abdominal segments ; cremaster not 
as described above, 
c. Entire body surface with coarse, deep punctures ; cephalic mar- 
gin of the movable abdominal segments with large lunate 
punctures and a distinct ridge with a row of large, very dis- 
tinct punctures just caudad of the ridge; cremaster short, 
rugose, slightly bifurcate, bearing six long hooked setae ; meso- 
thorax never with a deeply pitted caudal margin. 

Symmerista Hiibner. 
cc. Body usually punctate on the abdomen, but not on the appen- 
dages; movable abdominal segments sometimes with a slight 
ridge along the cephalic margin, but never with a distinct 
row of large punctures caudad of the ridge ; cremaster bifur- 
cate, but never with hooked setae ; mesothorax with a row of 
deep pits along its caudal margin with smooth quadrangular 
areas between, and partly covering them, 
d. First abdominal segment with a small tubercle on each side of 
the meson at the cephalic margin of the segment ; entire dor- 
sal surface of the tenth segment distinctly elevated and very 
rugose ; points of the cremaster divergent. 

Hyparpax Hiibner. 

dd. First abdominal segment without tubercles; entire dorsal 

surface of the tenth segment not elevated and rugose. 

e. Wings always touching on the meson; maxillae never as 

long as the wings ; cephalic end of body sometimes with 

two sharp, heavily chitinized spinous projections. 

Schizura Doubleday. 



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134 

ee. Wings adjacent on the meson but not touching; maxillae 
usually as long as the wings ; cephalic end of body without 
heavily chitinized spinous projections. 

Heterocampa Doubleday. 
The following species were examined : 
Melalophinae 

Melalopha inclusa Hiibner, apicalis Walker, albosigma Fitch 
Apatelodinae 

Apatelodes torrefacta Smith and Abbot, angelica Grote 
Cerurinae 

Harpyia borealis Boisduval 
Pygaerinae 

Datana mimstra Drury, nwdesta Beutenmiiller, angusii Grote and 
Robinson, chiriquensis Dyar, contracta Walker, drexelii Hy. 
Edwards, integerrima Grote and Robinson, major Grote and 
Robinson, palmii Beutenmiiller, robusta Strecker 
Notodontinae 

Symmerista albifrons Smith and Abbot 
Heterocampinae 

Hyparpax aurora Smith and Abbot 

Schizura ipotnoeae Doubleday, concinna Smith and Abbot, 

unicornis Smith and Abbot 
Heterocampa guttivitta Walker, bilineata Packard 

Family Digptidae 

The pupae of this family closely resemble those of the Geomet- 
ridae, but are more specialized than most of the genera in that fam- 
ily, although they show the epicranial suture (Fig. 115). The ap- 
pendages are arranged very much as in the Geometridae, but there is 
no trace of labial palpi, maxillary palpi, or prothoracic femora (Fig. 
114). The antennae are filiform, extending beyond the caudal mar- 
gin of the wings and about half way down on to the fifth abdominal 
segment. Each prothoracic leg extends cephalad between the sculp- 
tured eye-piece and the antenna. The distal ends of the prothoracic 
and mesothoracic legs and the maxillae are overlaid by the antennae, 
which lie adjacent on the meson at their distal ends. The abdomen is 
elevated at the dorso-meson to form a low ridge, and there are promi- 
nent hooked setae present on segments seven to ten as well as on the cre- 
master. This family contains a single American species, Phryganidia 
calif ornica Packard. The family has usually been placed between the 
Noctuidae and Notodontidae, and widely separated from the Geomet- 
ridae. The pupa shows no relationship to the noctuids, and is much 
more highly specialized than most members of that family. 



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SUPERFABilLY SPHINGOIDEA 



The members of this superf amily retain but one generalized char- 
acter, the presence of exposed portions of the prothoracic femora in 
some of the more generalized forms. The shape of the pupa is almost 
as distinctive as that of the larva, being usually fusiform, often with 
the head distinctly narrower than the thorax, giving the body a 
"shouldered" appearance. The epicranial suture is never present, the 
only distinct head suture remaining being that adjacent to the proxi- 
mal end of each antenna. The wings and maxillae are unusually long 
in most members of this superfamily and various means are taken to 
accomodate the extra length, particularly of the maxillae. The fourth 
abdominal segment is usually longer on the ventral surface than on 
the dorsal, and the wings are seldom broadly rounded at their caudal 
margins, but usually somewhat pointed. The position of the head is 
also changed in many species and found almost, or entirely, on the 
dorsal surface of the body. The mandibles are often very conspic- 
uous, being represented by strongly elevated tubercles. The protho- 
racic legs are usually about half the length of the wings and the meso- 
thoracic legs three fourths of their length. The metathoracic legs are 
seldom visible. The antennae are for the most part filiform and vary 
from two fifths to three fourths the length of the wings. In the genera 
Smerinthus, Paonias, Marumba, and Cressonia the antennae are con- 
siderably wider at their proximal end and slightly pectinate, being 
larger and longer in the male, and the whole appearance of the body 
reminds one strongly of the Satumiidae. These genera are in 
many respects the most specialized of the Sphingoidea, and some of 
them are found in cocoons. It is an interesting fact that the most spe- 
cialized forms in nearly all of the subfamilies of Sphingidae exam- 
ined, show relationship to the Satumiidae. This group is therefore 
considered as related to the Saturnioidea but more generalized. Cer- 
tain of its members resemble in some respects the Pyralididae and 
Gelechiidae. A cremaster is always present, usually triangular in out- 
line and often slightly bifurcate at the distal end. The abdomen often 
shows three or four transverse depressions on each segment which cor- 
respond to the annulet-like rings on the body of the larva. Except in 
rare instances the scar of the caudal horn of the larva is visible on the 
dorsum of the eighth abdominal segment. 

This superfamily contains a single family, the Sphingidae. For the 
most part the genera are easily distinguished, but there were no char- 
acters found that served to separate the genera Smerinthus and 
Paonias. The generic names of Dyar's list have been used as far as 
possible. A monograph of this family, giving tables and descriptions 



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136 

for the identification of subfamilies, genera, and species, has been pre- 
pared and will be published at an early date. The following table to 
genera is given simply for the identification of specimens and does not 
indicate natural relationships. The genera may be separated as fol- 
lows: 

a. Spiraeular furrows present on all the movable segments, that is, on 
abdominal segments 5—7. 

b. Maxillae with a free portion or so-called ** raised tongue-case" 
present — ^the maxillary loop, 
c. MaxiDary loop with its distal half recurved and touching the 
proximal portion, forming a prominent loop on the ventral sur- 
face of the body Herse Oken. 

cc. Maxillary loop never with the distal half recurved, the distal end 
touching the ventral surface of the body, 
d. Maxillary loop strongly arched from the ventral surface of the 
body, the greatest width of the sp^ce between the free por- 
tion of the maxilla and the ventral surface of the body always 
greater than the width of the maxiUary loop in that region, 
e. Dorsum of body with a strongly rugose triangular area on 
each side of the meson of the metathorax and a similar 
rugose band on abdominal segments 2-8 extending over the 
cephalic third of the segment ; lateral margins of the max- 
illary loop with a serrate appearance ; the distal end of the 
loop swollen and bulb-like and at least twice the width of 

the remaining portion Cocytius Hiibner. 

ee. Dorsum of body with a strongly elevated transverse ridge 
on each side of the meson on the metathorax, the ab- 
dominal segments punctate, but never rugose on the ce- 
phalic third ; lateral margins of the maxillary loop never 
serrate in appearance; the distal end of the loop some- 
what swollen but never twice the width of the remaining 

portion Phlegethontius Hiibner. 

dd. Maxillary loop never strongly arched, but usually closely 
applied to the ventral surface of the body, the space be- 
tween the loop and the ventral surface of the body never so 
great as the width of the loop in that region, 
e. Cephalic margin of abdominal segments 5-7 with one deep 
pocket-like furrow over each spiracle, 
f . Maxillary loop extending as far caudad as the distal ends 
of the prothoracic legs and occasionally beyond them. 

AtreiLs Grote. 
flf. Maxillary loop never extending as far caudad as the dis- 
tal ends of the prothoracic legs Dolba Walker. 

ee. Cephalic margin of abdominal segments 5-7 with two fur- 
rows, the ectal furrow shallow, the ental one deep and 
pocket-like. 



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137 

f. Spiracular furrows 5 mm. or more in transverse length 
and extending ventrad of the spiracle for a distance equal 

to the length of the spiracle CJilaenogramma Smith. 

flf. Spiracular furrows always less than 5 mm. in transverse 
length, seldom extending ventrad of the spiracle, if so, 
then for a distance less than the length of the spiracle. 

Sphinx Linnaeus, 
bb. MaxiUae of the usual type without a maxillary loop or so-called 
** raised tongue-case." 
c. Cephalic margin of abdominal segments 5-7 always with one 
deep pocket-like furrow over each spiracle, with or without a 
shallow ectal one. 
d. With one deep pocket-like furrow over each spiracle on abdom- 
inal segments 5-7. 
e. Surface of body spinose ; cremaster broad and truncate, the 
caudo-lateral angles usually produced into sharp points; 
caudal abdominal segments flattened on the ventral sur- • 
face, and with distinctly carinate lateral margins. 

Cressonia Qrote and Robinson, 
ee. Surface of body never spinose; cremaster pointed, trian- 
gular in outline, caudal abdominal segments never flat- 
tened on the ventral surface, nor with distinctly carinate 
lateral margins, 
f. Maxillae normally reaching to the caudal margin of the 
wings, slightly less in some individuals; mesothoracic 
wings never meeting on the meson caudad of the max- 
illae ; scar of the caudal horn never present on the dor- 
sum of the eighth abdominal segment. . :Lapara Walker, 
flf. Maxillae never more than five sevenths the length of the 
wings ; mesothoracic wings always meeting on the meson 
caudad of the maxillae; scar of the caudal horn always 
present on the dorsum of the eighth abdominal segment. 

Daremma Qrote. 
dd. With one deep pocket-like ental furrow and a shallower ectal 
one ; maxillae about two thirds the length of the wings. 

Ceratomia Harris, 
cc. Cephalic margin of abdominal segments 5-7 with three or four 
more or less interrupted furrows over each spiracle, the sur- 
face of the furrows often punctate like the remainder of the 
cephalic margin, 
d. Maxillae never half the length of the wings; average length 
of maxillae at meson 5-6 mm., sometimes 7 mm. in large speci- 
mens; dorsal cephalic margin of abdominal segments 5-7 
deeply punctate, the punctures adjacent to each other, giving 
it a honeycombed appearance, the cephalic margin separated 



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138 

from the remainder of the segment by a distinct carinate 
ridge. {Smerinthus Latreille. 

(Paonias Hiibner. 
dd. Maxillae half the length of the wings; average length of 
maxillae at meson 10 mm., sometimes exceeding this length ; 
dorsal cephalic margin of abdominal segments 5-7 punc- 
tate, the punctures somewhat scattered and not presenting 
a honeycombed appearance; cephalic margin never sep- 
arated from the remainder of the segment by a distinct 

ridge Marumba Moore. 

aa. Spiracular furrows never present on all of the movable segments, 
b. Spiracular furrows present on either one or two of the movable 
segments, 
c. Cephalic margin of abdominal segments 5 and 6 with three or 
four furrows over each spiracle, the furrows separated by 
sharply carinate ridges and extending almost to the meson on 
both dorsal and ventral surfaces; cremaster sparsely covered 
with short curved spines on the dorsal and lateral aspects. 

Hemaris Dalman. 
cc. Cephalic margin of fifth abdominal segment Avith three or more 
shallow furrows over each spiracle, which never extend on to 
the dorsal and ventral surfaces; cremaster never with short 
curved spines on any portion; labrum always on the dorsal 
surface of the head, 
d. Prothoracic femora apparent, 
e. Mandibular area always elevated and usually forming prom- 
inent tubercles ; cephalic margin of fifth abdominal segment 
with one ental furrow over each spiracle, extending for the 
^ entire width of the lateral aspect, and five or six interrupted 

ectal furrows with interrupted carinate ridges between; 
body surface dull; color light brown or coflfee-color with 

darker markings DeilephUa Ochsenheimer. 

ee. Mandibular area never elevated; cephalic margin of fifth 
abdominal segment with either three or four entire fur- 
rows over each spiracle; body surface highly polished; 
color black with red markings ; abdominal segments finely 

punctate on the cephalic half Dilophonota Burmeister. 

dd. Prothoracic femora never apparent ; cephalic margin of fifth 
abdominal segment with six or more interrupted furrows 
with short, more or less wavy interrupted ridges between; 
cephalic portion of maxillae slightly excurved and almost 
carinate on the meson; abdominal segments punctate over 

the entire length Theretra Hiibner. 

bb. Spiracular furrows never present on any of the movable abdom- 
inal segments, 
c. Pupae always less than two inches in length ; labrum usually at 
the cephalic end of the body but never on the dorsal surface. 



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d. Pupae with a prominent tubercle on the face-parts, just mesad 
of each glazed eye, and a prominent tubercle on the labrum ; 
maxillae with the proximal portion excurved and carinate on 

the meson Deidamia Clemens. 

dd. Pupae never with prominent tubercles on the face-parts or 

labrum ; maxillae never with the proximal portion excurved 

and carinate on the meson. 

e. Abdominal segments 5-7 with one or more interrupted rows 

of spines along the cephalic margin of the segment, more 

prominent on the dorsal surface Darapsa Walker. 

ee. Abdominal segments 5-7 never with spines along the ce- 
phalic margin of the segments, 
f . Mandibular area with prominent tubercles, 
g. Body surface rough, deeply punctate over the entire sur- 
face of the abdominal segments, especially segments 
8-10; body tapering rapidly from the fourth abdom- 
inal segment to the long pointed cremaster ; femora of 

prothoracic legs never apparent AmpJiion Hiibner. 

gg. Body surface smooth and polished, the cephalic portion 
of the segment with punctures of medium size, the 
caudal portion finely, sparsely punctate ; body taper- 
ing gradually from the fourth abdominal segment to 
cremaster; prothoracic femora apparent. 

Lepisesia Grote. 
flf. Mandibular area never elevated or with prominent tuber- 
cles, 
g. Dorsum of eighth abdominal segment with a row of 
large deep punctures or pits along the cephalic mar- 
gin; cremaster slender, never more than 2 mm. in 
length; body always light colored with darker mark- 
ings Ampelophaga Bremer and Grey. 

gg. Dorsum of eighth abdominal segment nev6r with a row 
of large punctures or pits along the cephalic margin ; 
cremaster always more than 2 mm. in length, the 
breadth equal to the length ; body always dark brown. 

Sphecodina Blanchard. 

cc. Pupae always more than two inches in length ; labrum either at 

the cephalic end or on the dorsal surface of the body. 

d. Color black, often marked with red ; body surface smooth and 

polished, with a very few small punctures on the abdominal 

segments; labrum at the cephalic end of the body. 

Pseiidosphinx Burmeister. 

dd. Color uniform dark brown; body surface roughened and 

densely, coarsely punctate ; labrum on the dorsal surface of 

the body, at least a distance equal to its own length away 

from the cephalic end of the body Pholus Hiibner. 



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140 

The following species were examined: 
Herse cingulata Fabricius 
Cocytius antaeus Drury 

Phlegethontius quinquemaculata Haworth, sexta Johannsen 
Atreus plebeia Fabricius 
Dolba hylaeus Drury 
Chlaeno gramma jasniinearum Boisduval. 
Sphinx kalmiae Smith and Abbot, drupiferarum Smith and Abbot, 

gordius StoU, luciHosa Clemens, chersis Hiibner, eremitus Hiib- 

ner 
Cressonia juglandis Smith and Abbot 
Lapara bombycoides Walker, coniferarum Smith and Abbot 
Daremmra catalpae Boisduval 
Ceratomia amyntor Geyer, undulosa Walker 
Smerinthus jammcensis Drury, cerysii Kirby 
Marumba modesta Harris 
Hemaris diffinis Boisduval, -gracilis Grote and Robinson, thysbe 

Fabricius 
Deilephila lineata Fabricius 
Dilophonota ello Linnaeus, alope Drury 
Theretra tersa Linnaeus 
Deidamia inscriptum Harris 
Darapsa pholus Cramer 
Amphion nessus Cramer 

Lepisesia gaurae Smith and Abbot, juanita Strecker 
Ampelophaga myron Cramer, versicolor Harris 
Sphecodina abbotti Swainson 
Pseudosphinx tetrio Linnaeus 
Pholus pandorus Hiibner, achemon Drury 

SuPERFAMiLY SATURNIOIDEA 

The pupae of this superf amily retain none of the generalized char- 
acters found in the families previously discussed and all the sutures 
of the head are obliterated, even those adjacent to the proximal ends 
of each antenna. The body is usually heavily chitinized, and although 
there are always a few setae present they are rarely visible to the un- 
aided eye. The superf amily is distinguished by the presence of broadly 
pectinate antennae, in the Ceratocampidae for about one third of the 
length, while in the Hemileucidae and Saturniidae they are broadly 
pectinate to the distal end, and generally have the stem of the flagel- 
lum elevated. The greatest width of each antenna is at least one fifth 



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of its length, often much wider, and the antennae seldom extend 
farther caudad than the prothoracic legs. There is a marked differ- 
ence in the sexes, the antennae of the male being much broader, some- 
what longer, and often meet on the meson, covering nearly all of the 
appendages except the wings. The legs are shorter than in most super- 
families, the prothoracic and mesothoracic legs usually either meeting 
or lying adjacent on the meson. The maxillae never reach the caudal 
margin of the wings, and their greatest length is not more than one 
third the length of the wings, but they are usually much shorter. The 
mesothoracic wings always lie adjacent on the meson and the meta- 
thoracic wings are often visible on the meson in the Saturniidae. The 
family Ceratocampidae has a row of broad triangular spines set on 
the edge of a flanged plate along both cephalic and caudal margins of 
the movable abdominal segments. They usually possess very long 
cremasters, which are always bifurcate at the distal end. The Hemi- 
leucidae have short cremasters, while there are none present in the 
Saturniidae and few of the genera have spines at the caudal end of 
the body. 

A paper on this superfamily, giving tables for the identification of 
families, genera, and species has been prepared and the first part, 
''The Classification of the Pupae of the Ceratocampidae and Hemi- 
leucidae," was published in the Annals of the Entomological Society 
of America, Vol. 7, 1914, pp. 277-300. The manuscript for the re- 
mainder is now in the hands of the editor of the same journal. The 
following tables of families and genera and the descriptions, with some 
slight additions and corrections, are taken from the paper mentioned 
above. The generic names in Dyar's list are used, but the genera are 
not arranged to indicate natural relationships. The superfamily 
Satumioidea may be divided into three families as follows : 

a. Pupae with the movable segments provided with flange-like plates 
which prevent their being telescoped, their lateral margins distinctly 
tapering caudad and each segment noticeably smaller than the seg- 
ment eephalad of it ; wings never elevated dorsad above the surface 
of the body ; a distinct cremaster always present ; stem of the flagel- 
lum of the antenna never elevated and distinct, 
b. Pupae with a distinctly bifurcate cremaster ; body usually rough- 
ened with spines on the exposed surface of the thorax and abdo- 
men; metathorax with prominent oblong tubercles on each side 
of the meson extending one third or more of the distance between 
the meson and the margin of the first pair of wings ; pupae always 

found in the ground .Ceratocampidae. 

bb. Pupae without a distinctly bifurcate cremaster; .body never 
roughened with spines on the exposed surface of the thorax and 



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abdomen; metathorax never with prominent oblong tubercles; 

pupae found either in cocoons or in the ground. . Hemileucidae. 
aa. Pupae with the movable segments never provided with flange-like 
plates which prevent their being telescoped, the lateral margins ap- 
proximately parallel so that the segments appear of equal size and 
are usually telescoped so that only the caudal margins of the seg- 
ments are visible; wings prominently elevated dorsad above the 
level of the body, the caudal portion of the mesonotum and meta- 
notum always depressed adjacent to the wings ; a distinct cremaster 
rarely present; stem of the flagellum of the antenna always ele- 
vated and distinct Saturnhdae. 

Family Hemileucidae 

Margins of the free segments never with a row of spines; the 
body surface never roughened with spines ; antennae with the stem of 
the flagellum never distinct, the central axis never set with spines, the 
antennae tapering gradually from the part having .the greatest width; 
maxillae never more than one sixth the length of the wings; proleg 
scars seldom prominent on abdominal segments five and six and rarely 
with the anal proleg scars visible; mesothoracic wings with the anal 
angles broadly rounded, usually at the cephalic margin of fourth ab- 
dominal segment, and usually reaching the caudal margin of the fourth 
abdominal segment ventrally ; metathoracic wings never produced be- 
low the anal angles of the first pair of wings and never visible in 
ventral view ; metathorax never with prominent tubercles ; abdominal 
segments 5 to 7 with the cephalic margin produced into a thick oblique 
flange-like plate directed caudad; cremaster short, never bifurcate 
at tip. 

Although not usually included with the Hemileucidae the genus 
Automeris is placed in this group owing to the very evident relation 
of the pupae to those of the genera Hemileuca and Pseudohazis. 
Morphologically they seem to be more nearly related to the Hem- 
ileucidae, but they are found in cocoons like the Saturniidae. Some 
of the members of this family pupate in the ground. 

The description of this family is of necessity very incomplete 
owing to lack of material. According to our available knowledge of 
the subject the three genera may be separated as follows : 

a. Cremaster bearing setae arranged in a tranverse row and spreading 

out fan-like Pseudohazis Grote and Robinson. 

aa. Cremaster never with setae, either with curved spines or without 
spines or setae of any kind, 
b. Cephalic part of segment above the flange-like plate either smooth 
or with fine longitudinal striations; pupae found in ground. 

Hemileuca Walker. 



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bb. Cephalic part of segment above the flange-like plate with sharp 
transverse ridges, deep furrows between; pupae found in 
cocoons Automeris Hiibner. 

The following species were examined : 
Pseudohazis eglanterina Boisduval 
Hemileuca tnaia Drury, burnsi Watson, olivae Cockerell 
Automeris pamina Neumoegen, io Fabricius, leucana Hiibner, incar- 
nata Walker 

Family CERATOCAMPmAE 

Body with the margins of the free abdominal segments usually 
bearing a row of spines, and the exposed surface of the tliorax and 
abdomen usually roughened with spines ; antennae never broadly pec- 
tinate throughout, but broadly pectinate and almost parallel for about 
one half the length, then narrowed rapidly to about half the greatest 
width, tapering gradually to a pointed tip, the stem of the flagelltun 
never distinct, the surface convex and the central axis of the antenna 
usually bearing one or two rows of small spines; maxillae never less 
than one fourth the length of the wings ; tips of the mesothoracic tarsi 
meeting obliquely on the meson, never lying adjacent on the meson; 
proleg scars very prominent on abdominal segments five and six, the 
scars for the anal prolegs often very conspicuous ; mesothoracic wings 
with the anal angles broadly rounded, usually located at the cephalic 
margin of the fourth abdominal segment and never reaching ventrad 
to the caudal margin of the fourth segment ; metathoracic wings never 
produced below anal angle of the mesothoracic wing and never visible 
in ventral view ; metathorax with distinct tubercles, more or less ob- 
long in outline, on each side the meson and extending more than one 
third the distance from the meson to the margin of the wing; the 
suture between the seventh and eighth segments never deep, or with 
distinct crenulations on its margins ; cremaster always present, usually 
long and bifurcate at tip. Five genera of this family have been 
described. One genus, Syssphinx, consisting of three species, was not 
available for study. The pupae of this family are always found in the 
ground. The remaining genera of Ceratocampidae can be separated 
by the following table : 

a. Surface of pupa never spinose ; cremaster broader than long, broadly 
and shallowly bifurcate, never over 2 mm. in length. . 

Citheronia Hiibner. 
aa. Surface of pupa spinose ; cremaster at least twice as long as broad, 
bifurcate at tip, always more than 2 mm. in length. 



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b. Thorax rugose with short isolated spines, abdominal segments 
not spinose, but bearing a row of spines along both cephalic and 
caudal margins of segments 1-7, the spines along the caudal 
margin of segments 5-7 much longer than the spines of the 

cephalic row BasUona Boisduval. 

bb. Thorax and abdominal segments densely spinose ; abdominal seg- 
ments 1-7 with a row of spines along both cephalic and caudal 
margins, the spines in the cephalic row on abdominal segments 
5-7 usually much longer than the spines in the caudal row ; 
maxillae, measured on meson, one fourth the length of the wings, 
c. Usually with prominent scattered spines on the thoracic segments, 
at least four times as long as those covering the segments ; an- 
tennae with the central axis bearing a row of prominent spines 
curved caudad ; if without prominent spines on the thoracic seg- 
ments and antennae, then the maxillae are one third the length 
of the wings, 
d. Eighth abdominal segment never with a prominent transverse 
ridge in the middle of the segment bearing a row of spines ; 
glazed eye-piece always lighter in color than the remaining 
surface of the body; species always more than an inch in 

length , Adelocephala Herrich-Schaeffer. 

dd. Eighth abdominal segment always with a prominent trans- 
verse ridge in the middle of the segment bearing a row 
of spines ; glazed eye-piece always the same color as the re- 
maining surface of the body; species never more than an 

inch in length Dryocampa Harris. 

cc. Without prominent scattered spines on the thoracic segments, 
the longest never four times the length of those covering the 
segments ; antennae with the central axis never bearing prom- 
inent spines, the spines never curving caudad ; maxillae always 
one fourth the length of the wings Anisota Hiibner. 

The following species were examined : 
atheroma regalis Fabricius 
Basilona imperialis Drury 
Adelocephala bicolor Harris, bisecta Lintner 
Dryocampa rubicunda Fabricius 

Anisota zirginiensis Drury, stigma Fabricius, senatoria Smith and 
Abbot, skinneri Biederman, constUaris Dyar 

Family Saturnudae 

The members of this family have the antennae broadly pectinate 
throughout, or nearly so, and the stem of the flagellum is usually dis- 
tinct and raised above the level of the pectinations. The maxillae are 
always short, never more than one third the length of the wings, and 



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usually very much shorter. The tibiae and tarsi of the prothoracic 
legs, and the tarsi of the mesothoracic legs lie adjacent on the meson, 
but never meet obliquely on the meson as they do in the ceratocampids. 
The mesothoracic wings always have their anal angles broadly rounded 
and the wings always reach the caudal margin of the fourth abdominal 
segment on the ventral surface. The metathoracic wings are pro- 
duced around the anal angles of the mesothoracic wings and usually 
form prominent angles on the fourth abdominal segment. The meta- 
thoracic wings always extend for at least a short distance along the 
caudal margin of the mesothoracic wings on the ventral surface of the 
body. The metathorax never has distinct oblong tubercles which are 
one third or more the width of the segment, such as are found in cera- 
tocampids. The suture between the seventh and eighth abdominal 
segments is never deep, with distinct crenulations on its margins, and 
is indistinct in many species. The cremaster, if present, is very short 
and is never bifurcate at the distal end. 

The known pupae of members of this family are found in silken 
cocoons. Some of these are very thick and tough, others thin and pa- 
pery. Only nine genera of this family have been available for study. 
These may be separated by the following table : 

a. Lateral margins of abdominal segments 5-7 never approximately par- 
allel, but tapering from the cephalic margin of the fifth segment, 
the lateral margins usually distinctly convex; caudal end of body 
usually with stout curved spines. 

b. Tenth abdominal segment never flattened into a transverse plate 

with the caudo-lateral angles produced into short lobes. 

c. Caudal end of body without spines; body surface with slightly 

wavy, transverse ridges with distinct furrows between; meso- 

thorax never with a prominent tubercle at the base of each wing. 

Copaxa Walker, 
ce. Caudal end of body with stout curved spines ; body surface never 
with slightly wavy, transverse ridges with distinct furrows be- 
tween; mesothorax with a prominent tubercle at the base of 
each wing, 
d. Lateral aspects of the cephalic margin of abdominal segments 
never with spiracular furrows; caudal end of body with an 
oval area set with slightly curved spines arranged in two 

groups and nearly all pointing outwards Telea Hiibner. 

dd. Lateral aspects of the cephalic margin of abdominal seg- 
ments 5-7 with spiracular furrows separated by slightly 
wavy carinate ridges; caudal end of body deeply rugose, 
with a slight concavity containing a circular group of 
strongly recurved spines Tropaea Hiibner. 



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bb. Tenth abdominal segment, viewed dorsally, in the form of a 
transverse plate, concave on the caudal margin, the caudo- 
lateral angles produced into lobes ; the other segments strongly 
concave in ventral view, with five short curved spines inserted 
close together in the caudo-lateral margin of each lobe. 

Agapema Neumoegen and Dyar. 
aa. Lateral margins of abdominal segments 5-7 always approximately 
parallel, the caudal end of the body never with stout curved spines, 
b. Maxillae always one fourth, or less, the length of the wings, the 
proximal two thirds of their margins never strongly concave; 
mesothoracic wings with their anal angles on the cephalic mar- 
gin of the fourth abdominal segment or caudad of that portion of 
the segment, 
c. Maxillae less than one fifth the length of the wings ; antennae of 
males with the sides tapering gradually to a pointed tip. 
d. Both eye-pieces never visible in either sex; glazed eye-piece 
seldom visible in the females, never in the males ; caudal end 
of abdomen never with a band of coarse setae, 
e. Eye-pieces never visible in either sex; caudal end of abdo- 
men never with spines or setae Callosamia Packard. 

ee. Glazed eye-piece visible in the females ; caudal end of abdo- 
men with a few very short sharp spines. 

Eupackardia Cockerell. 
dd. Both eye-pieces visible in either sex ; caudal end of abdomen 
with a transvense band of coarse setae. .Rothschildia Grote. 
cc. Maxillae never less than one fifth the length of the wings ; an- 
tennae of males with the sides approximately parallel for the 
greater part of their length, tapering rapidly to a blunt 
rounded tip ; a small portion of glazed eye-piece always visible 

in the females Samia Hiibner. 

bb. Maxillae always more than one. fourth the length of the wings ; 
the proximal two thirds of the lateral margins of the maxillae 
concave ; first pair of wings with their anal angles on the third 
abdominal segment opposite the second pair of abdominal 
spiracles Philosamia Grote. 

The following species were examined : 
Copaxa lavendera Westwood 
Telea polyphemus Cramer 
Tropaea luna Linnaeus 
Agapema galbina Clemens 

Callosamia promethea Drury, angulifera Walker 
Uupackardia calleta Westwood 
Rothschildia orizaha Westwood, jorulla Westwood 
Samia cecropia Linnaeus, gloveri Strecker, Columbia Smith, rubra 

Behr 
Philosamia cynthia Drury 



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147 

PHYI.OGENY 

The characters used as a basis for determining the phylogeny of 
the order are primarily: (i) the number of movable segments; (2) 
the freedom of the appendages; (3) the number of sutures present 
in the head; (4) the relative length of the body segments; (5) the 
presence or absence of visible labial palpi and maxillary palpi ; (6) the 
presence of exposed portions of the prothoracic femora in special- 
ized pupae; and (7) the method of dehiscence. 

In the most generalized forms there is complete freedom of mo- 
tion possible between the head and thorax, and between all the seg- 
ments of the thorax and abdomen with the exception of the eighth, 
ninth, and tenth abdominal segments, which are always fixed. As 
specialization proceeds, there is a gradual loss of motion ; first between 
the head and thorax, then between the segments of the thorax, and 
last of all between the different segments of the abdomen. The loss 
of motion in the abdomen begins first at the cephalic end, but by the 
time that complete motion of the second segment has been lost there 
begins a loss of motion of the seventh segment. This takes place first 
in the female, and there is a large series of forms, including the super- 
families Gracilarioidea, Tortricoidea, and Aegerioidea, which retain 
freedom of motion in the seventh segment of the male, while there is 
taking place at the cephalic end of the abdomen the loss of motion of 
the third abdominal segment. There are, however, a few gen- 
era of Gracilarioidea which have lost freedom of motion of all 
the body segments and which form the most specialized end of 
that series. The pupae which have lost motion of all the abdominal 
segments except the fourth, fifth, and sixth, are those usually referred 
to as obtected pupae. There are few pupae more specialized than 
those of the superfamily Gracilarioidea which retain freedom of 
motion of the seventh abdominal segment in the male, but there are 
a few generalized forms both in the Pyralidoidea and Papilionoidea in 
which this is the case, as it is also in the family Epermeniidae of the 
Yponomeutoidea. These three superfamilies are usually considered 
as more specialized than the Gracilarioidea. As the number of mov- 
able segments determines the position of a superfamily in the series 
it is readily seen that these superfamilies must be considered as more 
generalized than those in which motion is lost in the seventh segment 
in the male. It will be remembered that a segment is spoken of as 
movable when motion is possible between its caudal margin and the 
segment caudad of it. As the appendages become soldered to the body 
wall on the ventral surface no motion of this part of the segment is 
possible if the incision between its caudal margin and the next seg- 



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148 

nient is covered by the wings, therefore it can not be considered as a 
free segment. In many cases, however, dorsal movement of such seg- 
nients is possible, which gives the segment freedom of movement in 
certain directions; as, for instance, in curving the caudal end of the 
body cephalad on the ventral surface, well illustrated in the movements 
of most tortricids. Such forms must be considered as more general- 
ized than those which have lost entire motion of the segment, and thus 
the Pyralidoidea and Papilionoidea must occupy a lower position than 
the Yponomeutoidea, whose members have lost dorsal motion of the 
third abdominal segment, while the other two superf amilies mentioned 
retain it. There are certain specialized forms in other superfamilies 
in which motion is lost in all the body segments, notably in the family 
Elachistidae of the Gelechioidea and in certain genera of the family 
Nymphalidae in the Papilionoidea. There are also many genera in 
various families which retain movement in only one segment. 

The appendages of the generalized pupae are entirely free from 
^ach other and from the body wall and are often considerably spread 
out from the surface of the body so that the pupae strongly resemble 
those of the Trichoptera. In these forms there is but a slight degree 
of chitinization in any part of the body. The appendages are gprad- 
ually soldered down, however, first to each other, while all remain 
free from the body wall, and then there takes place a gradual sol- 
dering down of the appendages to the body wall, beginning first at 
the cephalic end of the abdomen. In many pupae the appendages are 
soldered to two, three, or four abdominal segments while the portion 
of the appendages caudad of these segments remains free and allows 
freedom of motion of the abdominal segments underneath. Such a 
<:ondition exists in many genera of the Aegerioidea and Gelechioidea. 
The pupae with free appendages could only exist successfully in pro- 
tected situations from which an easy egress was possible, and so they 
-are only found in cocoons, or in mines in leaves and stems of plants. 
Pupae with any other environment lost the freedom of the appendages 
much more rapidly, as in the case of the Lyonetiidae and some of the 
Papilionoidea. 

The number and arrangement of the sutures present on the head 
has already been discussed under the head of external morphology, 
pages 23 to 25. These sutures are gradually obliterated, beginning 
with the clypeo-labral, which is lost among very generalized pupae. 
The epicranial suture is one of the last to disappear, and its presence 
indicates the degree of specialization in many of the higher forms, as 
it is retained in some members of many superfamilies which are high 
in the series. 



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149 

The f ronto-clypeal suture is visible for a part of its length in most 
pupae, and is especially distinct for its entire length in some of the 
Celechioidea ; but dehiscence often showed the presence of this suture 
when it was impossible to locate it on the pupa. The part of this su- 
ture adjacent to the proximal end of each antenna is the last head su- 
ture to be obliterated, and it is lacking only in the Satumioidea. 

The segments of the body are more nearly of equal length in gen- 
eralized than in specialized forms, especially in the abdomen. The 
prothorax is short in the Micropterygoidea and becomes gradually 
longer in the specialized superfamilies. The metathorax is long in 
generalized forms and nearly equals the mesothorax in length. As 
specialization proceeds, the mesothorax becomes longer and the meta- 
thorax much shorter, so that the comparative length of these two seg- 
ments furnishes another means of determining the position of a super- 
family in the series. The abdominal segments also become consoli- 
dated, first at the caudal end of the body, where they gradually be- 
come shorter than the cephalic segments. After motion is lost in the 
cephalic segments, they, too, gradually shorten, until the movable seg- 
ments are much longer than any of the others. 

The presence of visible maxillary and labial palpi also furnishes 
an easy means for the identification of generalized forms. The labial 
palpi are retained throughout the series, but are gradually overlaid and 
concealed by the maxillae. The presence or absence of visible labial 
palpi, however, iadicates the degree to which specialization has pro- 
ceeded along a given line. Labial palpi are visible to some extent in 
some members of all superfamilies except the Saturnioidea. The 
maxillary palpi are usually the first to disappear, but these palpi are 
often present in the pupa, when lacking in the imago. The maxillary 
palpi in generalized forms reach the proximo-lateral angles of the 
maxillae, but gradually decrease in length until they are visible only 
as a small triangular area caudad of the sculptured eye-piece. 

When the appendages are free their position is considerably laterad 
■of that which they gradually assume as they become soldered to each 
other. The legs are folded in such a way that in generalized forms 
almost the entire femur of the prothoracic leg is exposed. Later the 
tibia and tarsus of this leg are folded so that their position is nearer 
the meson than formerly and the femur is entirely concealed. The 
presence of an exposed portion of the prothoracic femur is a general- 
ized condition which is retained by forms exceedingly specialized in 
other respects, and is found in some genera of Sphingidae. 

As to the method of dehiscence, there are several things to be 
moted, although all too little is known of this interesting phase of pupal 



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150 

life. There is a tendency for the generalized forms to emerge from 
the mine, cocoon, burrow, or other place of protection, as a" pupa, and 
consequently the body is provided with some structure which assists 
in its progress. The appendages and body segments are usually sep- 
arated from each other at dehiscence and the body splits along the 
median line of the vertex and thoracic segments, the vertex carrying 
the sculptured eye-pieces with it. The front, with the antennae, is com- 
pletely separated from the rest of the head parts in some forms, by a 
splitting along the epicranial suture on the dorsum, and along the 
f ronto-clypeal suture on the ventral surface. When the f ronto-clypeal 
suture is not entire it usually splits for a part of its length, thus al- 
lowing it to be considerably elevated. In specialized forms it is usually 
the imago which emerges, the pupal skin being left behind in the 
cocoon or other place of protection. The appendages and body seg- 
ments remain firmly soldered together and the imago escapes through 
the opening made by the splitting of the vertex, when present, the pro- 
thorax, and the mesothorax; or, if this is not sufficient, an irregular 
opening which does not follow the line of any suture is made in the 
cephalic end of the body. In these forms the eye-pieces remain at- 
tached to the other face-parts. 

The phylogeny of any group is usually determined by the devel- 
opment of a single character. Many workers have used the venation 
of the wings to arrange a series of genera or species in phylogenetic 
order. Others have used the genitalia, or the arrangement of setae. 
The pupae present many and varied characters which may be used to 
arrange such a series. In this investigation a series was arranged for 
each of the characters previously mentioned and the results of these 
series were combined. These characters have the advantage over 
those used by previous authors in that they comprise practically all of 
the important structures of the body and are all present in the same 
individual. It is quite probable that other characters might be used 
to indicate the development of the order, such as the number and ar- 
rangement of the genital apertures, the form of the spiracles, and the 
arrangement of setae, none of which have been investigated sufficiently 
to admit of their use in this paper. 

ACKNOWI.EDGMENTS 

The subject for this investigation was suggested by Professor A. 
D. MacGillivray in September, 1912. Since that time many species of 
Lepidoptera have been collected, both in the larval and pupal stage, 
and reared to maturity. The Graduate School of the University of 



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151 

Illinois has made liberal appropriations for the purchase of material, 
and a large series of named pupae has been obtained from the follow- 
ing sources : the American Entomological Company, Ward's Natural 
Science Establishment, the Kny-Scheerer Company, the New England 
Entomological Exchange, the New Jersey Entomological Company, 
and from Mr. Wm. Beutenmiiller, A. H. Manee, of Southern Pines, 
N. C, E. J. Oslar, of Denver, Colo., and Miss Annette F. Braun, of 
Cincinnati, Ohio. Dr. L. O. Howard, Chief of the Bureau of Ento- 
mology of the U. S. Department of Agriculture, Mr. August Busck, 
of the U. S. National Museum, and Dr. Wm. Barnes, of Decatur, 
111., have furnished a number of rare species which contributed in no 
small measure to the progress of the investigation. The collections of 
the Illinois State Laboratory of Natural History have been freely 
available, and for this courtesy Professor S. A. Forbes deserves our 
hearty thanks as well as his assistants, Mr. J. R. Malloch and Mr. C. 
A. Hart. Dr. T. H. McDunnough, of Decatur, 111., and Miss A. F. 
Braun have assisted in the identification of bred material, thus mak- 
ing available many species that otherwise could not have been used. 
The collections furnished by Miss Braun and Mr. Beutenmiiller 
deserve especial mention. Miss Braun collected, bred, and determined 
more than one hundred species of the so-called Microlopidoptera which 
really formed the working basis for this part of the investigation. 
Most of the work done on the Sphingidae has been from specimens 
furnished by Mr. Beutenmiiller, who has collected a large series of 
forms. Mr. Samuel Henshaw, of th^ Museum of Comparative Zool- 
ogy at Harvard, very kindly loaned a large series of Papilionoidea, 
most of which were collected by Dr. S. H. Scudder and described in 
his "Butterflies of the Northern United States and Canada." Profes- 
sor J. G. Needham, of the Department of Entomology at Cornell 
University, also loaned a large number of species from the collections 
of that University, which were especially valuable and could not be 
obtained elsewhere. 

It would be impossible in this brief space to enumerate the many 
ways in which Professor MacGillivray has assisted in the preparation 
of this paper. His interest in the work has been unfailing, and what- 
ever of value it may contain is due to his inspiration, encouragement, 
and helpful criticism. 



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BiBUOGRAPHY 

Chapman, T. A. 

'93. Some neglected points in the pupae of the heterocerous 
Lepidoptera. Trans. Ent. Soc. London, 1893:97-119. 

'93. On a lepidopterous pupa (Micropteryx purpurella) with 
functionally active mandibles. Trans. Ent. Soc. London, 
1893 * 255-265. 

'94. Some notes on the Microlepidoptera whose larvae are ex- 
ternal feeders. Trans. Ent. Soc. London, 1894:335-350. 

'96. Notes on pupae — Omeodes, Epermenia, Chrysocorys and 
Pterophorus. Trans. Ent. Soc. London, 1896: 129-147. 

Jackson, W. Hatchett. 

'91. Morphology of the Lepidoptera. Trans. Linn. Soc. London, 
Zool., Ser. 2, Vol. 5. 

Packard, A. S. 

'95. Attempt at a new classification of the Lepidoptera. Mono- 
graph of the Bombycine moths of America north of Mexico, 
Part L Memoirs of the National Academy of Sciences, 
7:56-83. 

Poulton, E. B. 

'91. The external morphology of the lepidopterous pupa. Trans. 
Linn. Soc. London, Zool., Ser. 2, 5 : 245-263. 

Scudder, S. H. 

'89. The butterflies of the Eastern United States and Canada. 3 
vols. 

Tutt, J. W. 

'00. A natural history of the British Lepidoptera, 2 : 38-100. 



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153 



Pirates 



The following plates show in outline the principal structures of 
pupae of many of the families discussed in this paper. No attempt 
has been made to show all of the setae, spines, or tubercles which may 
occur, but only those which are most important and are of taxonomic 
value. 

The following abbreviations have been used : 

genae 

glazed eye-piece 

genital opening 

labrum 

prothoracic leg 

meeothoracic leg 

metathoracic leg 

labial palpi 

mandibles 

maxillary palpus 

mesothorax 

meeothoracic spiracle 

metathoraz 

maxilla 

prothorax 

pilifer 

proleg scar 

spiracle 

sculptured eye-piece 

spiracular furrow 

tegulae 

tubercle scar 

vertex 

ventro-spiracular row of tubercles 

mesotiioracic wing 

metathoracic wing 



a, 


antennae 


gf 


al-alO, abdominal segments 1-10 


ge, 


af, 


alar furrow 


go, 


ao. 


anal opening 


lb, 


ar, 


anal rise 


li, 


at, 


invaginations for the anterior 


1^ 




arms of the tentorium 


1., 


d, 


dypeus 


Ip, 


Cl8, 


dypeo-labral suture 


md. 


cdm, 


caudal margin of an abdominal 


mp. 




segment 


ms, 


cm, 


cephalic margin of an abdominal 


msp, 




segment 


mt, 


cr, 


cremaster 


mx. 


cs. 


cremastral setae 


Vf 


cxl. 


coxa of the prothoracic leg 


Pf, 


cx2, 


coxa of the mesothoracic leg 


psc, 


cx3. 


coxa of the metathoracic leg 


8, 


dlt, 


dorso-lateral row of tubercles 


se. 


dmt. 


dorso-mesal row of tubercles 


sf. 


dst, 


dorso-spiracular row of tubercles 


t, 


es, 


epicranial suture 


ts, 


f, 


front 


▼, 


fcs. 


fronto-clypeal suture 


vst. 


fl 


femur of the prothoracic leg 


wl, 


'2, 


femur of the mesothoracic leg 


w2, 


^, 


flanged plate 





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INDEX TO GENERA AND HIGHER GROUPS 



Achatodes, 114. 

AcrobasiSy 75. 

AcrocercopSy 67. 

Acrolopliidae, 45, 46. 

Acronycta, 114, 115, 119. 

Acronyctinae, 108, 113. 

Adelocephala, 144. 

Aegeriidae, 24, 48, 49, 52. 

Aegerioidea, 27, 31, 48-51, 147, 148. 

Agapema, 146. 

Agaristinae, 109, 112. 

Aglais, 91. 

Agonopteryx, 104, 105. 

Agranlis, 92. 

Agratinae, 107, 109, 110. 

Agrotis, 109. 

AlsophUa, 127. 130. 

Alypia, 113. 

Ambljscirtes, 79, 81. 

Ampelophaga, 139. 

Amphion, 139. 

AnacampsiB, 102. 

Anaea, 89, 93, 94. 

Anaeinae, 89, 94. 

Ancjlis, 52. 

Aula, 129. 

Anisota, 144. 

Anomie, 107, 115. 

AnoBia, 94. 

Antispila, 63. 

Apantesis, 120. 

Apatelodes, 132, 133. 

Apatnrinae, 89, 93. 

Apaturini, 94. 

Archips, 29, 55, 57, 58. 

Arctudae, 107, 116, 119. 



Argynnini, 90, 91. 
ArgynniB, 92. 
Argyresthia, 98. 
Argyrotoxa, 56. 
AristoteUa, 101, 102, 103. 
Atreus, 136. 
Atteva, 71, 72. 
Attevidae, 69, 70, 71. 
Automeris, 143. 

B 
Balsa, 116, 117. 
Basilarehia, 93. 
Basilarehinae, 89, 93. 
Basilona, 144. 
Bedellia, 65. 
Bembeda, 49, 50. 
Bombycidae, 123, 124. 
Bombyooidea, 33, 96, 123125. 
Bombyx, 124. 
Brenthia, 47, 48, 63. 
Brenthis, 92. 
Brephos, 130. 
Baccnlatrigidae, 61, 64. 
Bucculatriz, 64. 
Butalis, 100. 


Caenurgia, 118. 
Callosamia, 146. 
Cissia, 95. 
Citberonia, 148. 
Cleora, 130. 
Gocceius, 82.' 
Cocytius, 136. 
Coleopbora, 98. 
Coleopboridae, 96, 98. 
Oopaxa, 145. 



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Ooptodisca, 63. 
Coptotriche, 63. 
Cosmopterjgidae, 99, 106. 
Cosmopteryx, 106. 
Cosmotriche, 124. 
Cossidae, 39, 40. 
Ck>88iiiae, 41. 

Cossoidea, 26, 31, 37, 38-41. 
Crambinae, 72, 73, 74. 
Cremastobombjeia, 68. 
Cressonia, 135, 137. 
Gtenucha, 119, 120. 
CuculUanae, 107, 108, 110. 
Oyaniris, 84. 
Cymatopbora, 130. 
Calpodes, 79, 81. 
Cameraria, 64, 67, 68, 69. 
Canarsia, 76. 
Carpocapsa, 52, 53. 
Oatocala, 117. 
Catocalinae, 109, 110, 117. 
Cenopis, 58. 

Ceratocampidae, 140, 141, 143. 
Ceratomia, 137. 
Cercyonis, 95. 
Charadra, 116, 119. 
Charidryas, 92. 
Chlaenogramma, 137. 
Chloridea, 112. 
Chlorippe, 93. 
Choreutis, 47, 48, 63. 
Chrysopeleia, 104. 
Chrysopeleiidae, 99, 104. 
Ghrysophanus, 83, 84. 
Oinclidia, 93. 
Cingilia, 129. 
Ginglis, 128. 
Cirphis, 110, IIL 

D 

Darapsa, 139. 
Daremma, 137. 
Dasychira, 122. 
Datana, 133. 
Deidamia, 139. 
Deilepbila, 138. 
Depressaria, 104, 105. 



Desmia, 77. 
Diacrisia, 120. 
Dilopbonota, 138. 
Dioptidae, 125, 126, 134. 
Dolba, 136. 
Dryocampa, 144. 

E 

Ecpantheria, 120. 

Ectropis, 130. 

Elachista, 65, 104, 106. 

Elachistidae, 62, 96, 98, 99, 100, 106. 

Enarmonia, 52, 54. 

Ennominae, 127. 

Ennomos, 129. 

Eoi8,128. 

Epagoge, 57, 58. 

Epargyrens, 82. 

Epermenia, 97. 

Epermeniidae, 59, 95, 96, 97, 147. 

Epbestia, 74, 75. 

Epiblemidae, 47, 52. 

Epinotia, 52, 53, 54. 

Epipaschiinae, 72, 73, 77. 

EpisimuB, 55. 

Erannis, 131. 

Eriocraniidae, 23, 24, 25, 26, 35, 37, 48, 
59, 60, 62. 

Eriopus, 110, 112. 

Estigmene, 120. 

Euchaetias, 119. 

Euclea, 43. 

Eucleidae, 24, 42, 43, 44. 

Eucleoidea, 25, 26, 29, 31, 37, 41-44, 62. 

Eucosma, 52, 53. 

Eudamus, 82. 

Eulonche, 114, 119. 

Eunetis, 117. 

Eupackardia, 146. 

Euparthenos, 118. 

Euphoeades, 86. 

Eupbydryas, 92. 

Euploeinae, 89, 94. 

Euproctis, 122. 

Euptoieta, 92. 

Eurema, 88. 

Eurymus, 88. 



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Euthisanotia, 118. 
Euvanessa, 91. 
Evippe, 102. 
Exartema, 55, 50. 



Feniseca, 78, 83, 8i. 

G 

Oalleria, 74. 

Galleiidae, 26. 

Gallerinae, 72, 73. 

Gelediia, 103. 

Gelechiidae, 99, 100, 101, 105, 135. 

Gelechioidea, 23, 33, 96, 98-106, 148, 149. 

Geometridae, 29, 125, 126, 134. 

Gnorimoschema, 101. 

Gracilaria, 26, 60, 66. 

Gracilariidae, 61, 62, 65, 69. 

Gracilariinae, 66. 

Gracilarioidea, 23, 26, 31, 58-69, 147. 

Graptolitha, 110. 

H 

Hadena, 110, 112. 
Hadeninae, 108, 110. 
Haematopsis, 126, 129. 
Hallsidota, 119. 
Hapalia, 110. 
Haploa, 120. 
Hannologa, 57. 
Harpyia, 133. 
Harrisiiia, 44. 
Heliodinidae, 45, 47, 63. 
Heliozelidae, 24, 59, 60, 62. 
Hemaris, 138. 
Hemerocampa, 121, 122. 
Hemileuca, 142. 
Hemileucidae, 140, 141, 142. 
Hemimene, 52, 54. 
Heodes, 83, 84. 
Hepialidae, 23, 24, 37, 38, 40. 
Hepialoidea, 30, 37-38, 39, 41. 
Herse, 136. 

Hesperiidae, 27, 78, 79, 80. 
Hesperioidea, 78. 



Heterocampa, 134. 
Homopyralis, 107, 114. 
Hydria, 128. 
Hjdriomemnae, 127. 
Hjparpaz, 133. 
Hjpeninae, 107, 109, 116. 
Hyphantria, 120. 
Hjpocolpus, 46. 
Hypsopygia, 74. 



Incisalia, 84. 
Iphiclides, 86. 
Isia, 120. 

J 

Junonia, 91. 

L 

Laertias, 86. 

Lagoa, 43. 

Lanthape, 78. 

Lapara, 137. 

Laphygma, 111. 

Lasiocampa, 124 

Lasiocampidae, 123, 124. 

Laverna, 100. 

Lavernidae, 99, 106. 

Lepisedia, 139. 

Leucauthiza, 67. 

Liparidae, 29, 107, 119, 121, 123. 

LithocoUetinae, 66, 67. 

Lithocolletis, 59, 67, 68. 

Lophoptilus, 100. 

Lycaenidae, 27, 78, 79, 83. 

Lycia, 131. 

Lycophotia, 110, 112. 

Lyonetiidae, 59, 60, 61, 64. 



Malacosoma, 124. 
Marmara, 67, 68. 
Marumba, 135, 138. 
Megalopygidae, 42, 43, 44. 
Megathymidae, 78, 79. 
Megathymus, 80. 
Melalopha, 132. 
Meliana, 111. 



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Melitaeinae, 90, 92. 

Mellisopus, 53. 

MemjthruSy 49, 50. 

Menesta, 105. 

Meroptera, 75. 

Micropterygoidea, 24, 30, 35-37, 149. 

Micropteryx, 20. 

Mineola, 75. 

Mitura, 84. 

Mnemonica, 35, 37. 

Mominae, 108, 116. 

Momphidae, 106. 

Monima, 110, 111. 

N 
Nacophora, 129. 
Nepticulidae, 26, 29, 41, 48, 59, 60, 61, 

65. 
Noctua, 110. 
Noctuidae, 107, 134. 
Noctuoidea, 26, 33, 96, 107-123. 
Notodontidae, 125, 132, 134. 
Notodontoidea, 34, 76, 96, 125-134. 
Nycteolidae, 119. 

Nymphalidae, 78, 79, 87, 88, 148. 
Nymphaliiiae, 89, 90, 93. 



Oecophoridae, 99, 104. 
Oeneinae, 90, 95. 
Oeneis, 78, 95. 
Oiketicufl, 40. 
Olene, 122. 
Olethreutes, 55, 56. 
Olethreutidae, 52, 54. 
Ornix, 59, 66. 
OxyptUus, 70, 71. 

P 

Paleacrita, 127. 

Pantagrapha, 77. 

Paonias, 135, 138. 

Papilio, 20, 86. 

Papilionidae, 78, 79/ 85, 87. 

Papilionoidea, 23, 24, 27, 32, 65, 69, 76, 

78-95, 96, 147, 148. 
Parectopa, 66. 



Parharmonia, 50. 
Peronea, 56, 57. 
Phaecasiophora, 57, 58. 
Pheocyma, 118. 
Philobia, 1^9. 
Philosamia, 146. 
Phlegethontius, 136. 
Phlyctaenia, 77. 
Pholisora, 82. 
Pholus, 139. 
Phryganidia, 134. 
Phthorimaea, 103. 
Phyciodee, 92. 
Phycitinae, 71, 73, 74, 75. 
Phyllocnistidae, 60, 61, 65, 
Phyllocnistis, 68. 
Physostegania, 130. 
Phytometra, 116. 
Phytometrinae, 108, 115. 
Pieridae, 78, 79, 87. 
Pinipestis, 75, 76. 
Plathypena, 116, 117. 
Platynota, 57, 58. 
Platyptilia, 70, 71. 
Plodia, 74, 75. 
Plusiodonta, 107, 114, 115. 
PluteUa, 97. 
Podosesia, 29, 50. 
Polia, 110, 112. 
Polychrosis, 55, 56. 
Polygonia, 91. 
Pontia, 88. 
Porthetria, 121, 122. 
Prionoxystus, 40, 41. 
Prodenia, IIL 
Prodoxidae, 44, 45. 
Prodoxus, 46. 
Proleucoptera, 65. 
Prolimacodes, 43. 
Protoparee, 20. 
Pseudanaphora, 46. 
Pseudohazis, 142. 
Pseudosphinx, 139. 
Psilocorsis, 104, 105. 
Psorosina, 76. 
Psychidae, 39, 40. 
Psychomorpha, 118. 



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159 



Pterophoridae, 28, 69, 70. 
Pterophorusy 70, 71. 
Pyralididae, 69; 70, 72, 185. 
PyraUdoidea, 24, 32, 69-78, 96, 147, 
Pyralinae, 73, 74. 
Pyralis, 74. 
Pyrausta, 76, 77. 
Pyraustidae, 23. 
Pyraustinae, 73, 76. 
Pyromorpbidae, 42, 44. 
Pyrrhia, 112. 



Recurvaria, 101, 102, 103. 
Bhodophora, 111. 
Rothschildia, 146. 
BusticuB, 84. 

S 
Sabulodes, 129. 
Samia, 146. 
Sanninoidea, 50. 
SaiTothripinae, 109, 118. 
Sarrothripus, 119. 
Saturniidae, 25, 28, 125, 135, 140, 

142, 144. 
Saturnioidea, 22, 34, 96, 123, 140- 

149. 
Satyiinae, 78, 90, 94, 95. 
Satyrodes, 95. 
Schiznra, 133. 
Scythridae, 96, 99, 100. 
ScythriB, lOL 
Sibine, 43. 
Sitotroga, 103. 
Smerinthus, 135, 138. 
Sparganothidae, 52, 57. 
Sphecodina, 139. 
Sphingidae, 23, 24, 25, 29, 135. 
Sphingoidea, 34, 96, 135-140. 
Sphinx, 137. 
Stenoma, 105. 
Stenomidae, 99, 104, 105. 
Sterrhinae, 127. 
Sthenopis, 38. 
Symmeiista, 133. 
Synanthedon, 50. 
Synehloe, 88. 



148. 



141, 
146, 



Syngrapha, 116. 
Syntomidae, 119. 
Syssphinx, 143. 



T 



Telea, 27, 145. 

Telphusa, 102. -. — 

Tephrodystis, 128. 

Thanaos, 82. 

Thecla, 84. 

Theretra, 138. 

Thiodia, 54. 

Tholeria, 77. 

Thorybes, 82. 

Thyridopteryx, 40. 

Tinea, 47. 

Tineidae, 45, 47. 

Tineoidea, 28, 31, 44-48, 59, 60. 

Tischeria, 63. 

Tischeriidae, 61, 63, 68. 

Tmetocera, 54. 

Tolype, 123, 124. 

Tortricidae, 52, 56. 

Tortricoidea, 28, 29, 31, 44, 45, 48, 49, 

51-58, 59, 147. 
Trichotaphe, 102. 
Tropaea, 27, 145. 
Trypanisma, 103. 

U 

Uranotes, 84. 
Utetheisa, 120. 

V 
Vanessa, 91. 
Vanessini, 90. 

X 
Xanthotype, 129. 
Xylorictidae, 105. 

Y 
Yponomeuta, 97. 
Yponomeutidae, 69, 71, 96, 97, 98, 100, 

106. 
Yponomeutoidea, 26, 32, 96-98, 147, 148. 
Ypsolophus, 102. 

Z 

Zale, 118. 
Zelleria, 97. 
Zeuzera, 41. 
Zeozerinae, 41. 



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Plate XIX 

Fig. 1. Mnemonica auricyanea, ventral view, male. 

Pig. 2. Mnemonica auricyanea, lateral view, female. 

Pig. 3. Mnemonica auricyanea, dorsal view, female. 

Pig. 4. Mnemonica auricyanea, dorsal view, caudal end of abdomen, 

male. 
Pig. 5. Mnemonica auricyanea, ventral view, caudal end of abdomen, 

male. 
Pig. 6. Mnemonica auricyanea, dorsal view, caudal end of abdomen, 

female. 
Pig. 7. Mnemonica auricyanea, ventral view, caudal end of abdomen, 

female. 
Pig. 8. Sthenopis thule, ventral view, male. 
Pig. 9. Sthenopis thule, lateral view, male. 
Pig. 10. Sthenopis thule, dorsal view, male. 
Pig. 11. Thyridopteryx ephemeraeformis, ventral view, male. 
Pig. 12. Thyridopteryx ephemeraeformis, lateral view, male. 
Pig. 13. Thyridopteryx ephemeraeformis, dorsal view, male. 
Pig. 14. Thyridopteryx ephemeraeformis, ventral view, female. 
Pig. 15. Prionoyxstus robiniae, face-parts, male. 
Fig. 16. Zeuzera pyrina, ventral view, male. 



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Platk XIX 




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Plate XX 

Pig. 17. Lagoa crispata, ventral view, female. 

Fig. 18. Lagoa crispata, dorsal view, female. 

Fig. 19. Euelea ehloris, ventral view, female. 

Fig. 20. Euelea ehloris, dorsal view, male. 

Fig. 21. Euelea ehloris, mesothoracic spiracle and adjacent parts. 

Fig. 22. Euelea ehloris, cephalic view of head and prothorax. 

Fig. 23. Prolimacodes scapha, ventral view, male. 

Pig. 24. Harrisina americana, ventral view, female. 

Fig. 25. Harrisina americana, dorsal view, female. 

Fig. 26. Prodoxus quinquepunctella, face-parts. 

Fig. 27. Prodoxus quinquepunctella, lateral view. 

Fig. 27a. Prodoxus quinquepunctella, tubercle of the eighth abdominal 

segment, lateral view. 
Fig. 28. Hypocolpus mortipennellus, ventral view, female. 
Fig. 29. Hypocolpus mortipennellus, dorsal view, female. 
Fig. 30. Tinea pellionella, ventral view, male. 
Fig. 31. Tinea pellionella, doi-sal view, male. 



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Plate XX 




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Plate XXI 

Fig. 32. Brenthia pavonacella, ventral view, female. 

Fig. 33. Brenthia pavonacella, dorsal view, male. 

Fig. 34. Choreutis gnaphiella, ventral view, female. 

Fig. 35. Choreutis gnaphiella, dorsal view, female. 

Fig. 36. Podosesia syringae, ventral view, female. 

Fig. 37. Synanthedon pietipes, dorsal view, head, thorax and abdominal 

segments 1-3. 
Fig. 37a. Synanthedon pietipes, dorsal view, abdominal segments 8-10. 
Fig. 38. Ancylis comptana, ventral view, male. 
Fig. 38a. Ancylis comptana, anal rise, lateral view. 
Fig. 39. Ancylis comptana, dorsal view, male. 
Fig. 40. Exartema ferriferanum, ventral view, male. 
Fig. 41. Peronea minuta, lateral view, male. 



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Plate XXI 




37 « 



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Plate XXII 

Fig. 42. Peronea ininuta, ventral view, female. 

Fig. 43. Peronea minuta, showing the eonneetion of the eye-pieees with 

the vertex after removal of antennae at dehiscenee. 
Fig. 44. Archips argyrospila, ventral view, female. 
Fig. 45. Graeilaria negundella, ventral view, male. 
Fig. 46. Graeilaria negundella, terminal segments of antenna. 
Fig. 47. Graeilaria sassafrasella, eephalie end of prothoraeie and meso- 

thoraeie legs, with adjoining area supposed to be the location 

of the maxillary palpus. 
Fig. 48. Nepticula platanella, ventral view, male. 
Fig. 49. Nepticula platanella, dorsal view, male. 
Fig. 50. Antispila cornifoliella, ventral view, male. 
Fig. 51. Coptotriche zelleriella, ventral view, male. 
Fig. 52. Coptotriche zelleriella, doi'sal view, male. 
Fig. 52a. Coptotriche zelleriella, lateral view, caudal end of abdomen. 
Fig. 52b. Coptotriche zelleriella, tips of strongly chitinized setae. 
Fig. 53. Tischeria malifoliella, doi*sal view, female. 



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Plate XXII 




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Plate XXIII 

Fig. 54. Tischeria heliopsisella, dorsal view, male. 
Fig. 54a. Tischeria heliopsisella, lateral view, caudal end of abdomen. 
Fig. 55. Bucculatrix sp., ventral view, male. 
Fig. 56. Bucculatrix sp., dorsal view, female. 
Fig. 56a. Bucculatrix sp., lateral view of head. 
Fig. 57. Bedellia somnulentella, ventral view. 
Fig. 58. Bedellia somnulentella, dorsal view. 
Fig. 59. Proleucoptera smilaciella, ventral view. 
Fig. 60. Proleucoptera smilaciella, dorsal view. 
Fig. 61. Cameraria hamadryadella, ventral view, male. 
Fig. 62. Cameraria hamadryadella, dorsal view, male. 
Fig. 62a. Cameraria hamadryadella, lateral view, head. 
Fig. 62b. Cameraria hamadryadella, dorsal view fifth and sixth abdom- 
inal segments, female. 
Fig. 63. Gracilaria negundella, dorsal view, male. 



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PI.ATE XXIII 




60 « 



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Pl.\te XXIV 

Fig. 64. Lithocolletis argentinotella, ventral view, female. 

Fig. 65. Lithocolletis argentinotella, dorsal \4e\v, male. 

Fig. 66a. Lithocolletis lucidicostella, dorsal view, cremaster. 

Fig. 66b. Lithocolletis tiliacella, dorsal view, cremaster. 

Fig. 67. Phyllocnistis insignis, ventral \new, male. 

Fig. 68. Eperminia pimpinella, ventral view, male. 

Fig. 69. Galleria melonella, lateral view, male. 

Fig. 70. Oxyptilus tenuidactj lis, ventral view. 

Fig. 71. Oxyptilus tenuidactylis, dorsal view. 

Fig. 72. Atteva aurea, ventral view, male. 

Fig. 73. Atteva aurea, dorsal view, male. 

Fig. 74. Ephestia kuehniella, ventral view, female. 

Fig. 75. Mineola indiginella, dorsal view. 



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Plate XXIV 




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Plate XXV 

g. 76. Pyrausta futilalis, ventral view, female. 

g. 77. Epargyreus tityrus, lateral view. 

g. 78. Calpodes ethlius, ventral view. 

g. 79. Cyaniris ladon, ventral view. 

g. 80. Oeneis semidea, ventral view. 

g. 81. Euvanessa antiopa, lateral view, fifth abdominal segment. 

g. 82. Zelleria celastrusella, ventral view, male. 

g. 83. Yponomeuta malinellus, ventral view. 

g. 84. Plutella maeulipennis, dorsal view. 

g. 85. Argyresthia f reyella, ventral view, female. 

g. 86. Argj^resthia f reyella, dorsal view, female. 

g. 87. Coleophora malivorella, ventral view. 

g. 88. Lophoptilus eloisella, ventral view, female. 

g. 88a. Lophoptilus eloisella, lateral view, eaudal end of abdomen. 

g. 89. Scythris eboraeensis, ventral view. 



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Plate XXV^ 




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Plate XXVI 

Fig. 90. Golechia serotinella, ventral view. 

Pig. 91. Trichtotaphe flavoeostella, dorsal view. 

Fig. 92. Trichtotaphe flavoeostella, ventral view. 

Fig. 93. Evippe prunifoliella, ventral view. 

Fig. 93a. Evippe prunifoliella, lateral view caudal end of abdomen. 

Fig. 94. Ypsolophus citrifoliella, ventral view. 

Fig. 95. Chrysopeleia ostryaeella, ventral view. 

Fig. 96. Stenoma schlaegeri, ventral view. 

Fig. 96a. Stenoma schlaegeri, dorsal and lateral views, fourth abdom- 
inal segment. 

Fig. 96b. Stenoma schlaegeri, tips of large lateral setae. 

Fig. 97. Psilocorsis quercicella, ventral view. 

Fig. 98. Psilocorsis quercicella, lateral view. 

Fig. 99. Cosmopteryx clandestinella, ventral view. 
Fig. 100. Elachista praelineata, ventral view. 



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Plate XXVI 




98 It 



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Plate XXVII 

Pig. 101. Agrotis biearnea, dorsal view. 

Pig. 102. Cirphis unipuneta, ventral view. 

Pig. 103. Eulonche oblinita, ventral view. 

Fig. 104. Isia Isabella, lateral view. 

Pig. 105. Hemerocampa leueostigma, ventral view. 

Pig. 106. Malacosoma disstria, ventral view. 

Pig. 107. Bombyx mori, ventral view. 

Pig. 108. Brephos infans, ventral view. 

Pig. 109. Hydria undulata, dorsal view, cremastral setae not present. 

Pig. 110. Cymatophora ribearia, dorsal view, caudal abdominal segments 
and eremaster. 

Pig. 111. Tephroclystis absinthiata, dorsal view, caudal abdominal seg- 
ments and eremaster. 

Pig. 112. Melalopha inclusa, dorsal view, abdominal segments and ere- 
master. 

Pig. 113. Datana angusii, dorsal view, caudal abdominal segments and 
eremaster. 

Pig. 114. Schizura ipomoeae, dorsal view, caudal abdominal segments and 
eremaster. 

Pig. 115. Phrjganidia calif ornica, ventral view. 

Pig. 116. Phryganidia calif ornica, dorsal view. 



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Plate XXVII 




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VITA 

The writer was born at Kempt Shore, Nova Scotia, and educated 
in the public schools of that province, graduating from the Provincial 
Normal School in 1902. For three years she held a position in one 
of the units of Sir William MacDonald's consolidated school garden 
system and then entered Cornell University, graduating in 1908 with 
the degree of Bachelor of Science in Agriculture. The next two years 
were spent as teacher of agriculture in the High School at Hampton, 
Virginia, and as supervisor of nature study and school gardening 
in the public schools of that city. A similar position was held in the 
public schools at Gary, Indiana, until 1912. The summers of 1909- 
191 1 were spent as instructor in agriculture and nature study in the 
State Normal Institute at Emory, Virginia. From September, 1912, 
until the present time the writer has been a graduate student at the 
University of Illinois, as scholar in Entomology during the year 1912- 
1913 and as fellow in 1914-1915. She received the degree of Master 
of Science in 19 13. 

The writer is a member of the Entomological Society of America 
and of the honorary society of Sigma Xi. She has published "A Class- 
ification of the Pupae of the Ceratocampidae and Hemileucidae," in 
the Annals of the Entomological Society of America for 1914, and 
"Homology of the Mouth Parts of the Preimago in the Lepidoptera," 
in the Journal of Entomology and Zoology for 1915- 



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